Tone Sequences Research Articles

Cued Memory Reactivation During SWS Abolishes the Beneficial Effect of Sleep on Abstraction.

Extracting regularities from stimuli in our environment and generalizing these to new situations are fundamental processes in human cognition. Sleep has been shown to enhance these processes, possibly by facilitating reactivation-triggered memory reorganization. Here, we assessed whether cued reactivation during slow wave sleep (SWS) promotes the beneficial effect of sleep on abstraction of statistical regularities. We used an auditory statistical learning task, in which the benefit of sleep has been firmly established. Participants were exposed to a probabilistically determined sequence of tones and subsequently tested for recognition of novel short sequences adhering to this same statistical pattern in both immediate and delayed recall sessions. In different groups, the exposure stream was replayed during SWS in the night between the recall sessions (SWS-replay group), in wake just before sleep (presleep replay group), or not at all (control group). Surprisingly, participants who received replay in sleep performed worse in the delayed recall session than the control and the presleep replay group. They also failed to show the association between SWS and task performance that has been observed in previous studies and was present in the controls. Importantly, sleep structure and sleep quality did not differ between groups, suggesting that replay during SWS did not impair sleep but rather disrupted or interfered with sleep-dependent mechanisms that underlie the extraction of the statistical pattern. These findings raise important questions about the scope of cued memory reactivation and the mechanisms that underlie sleep-related generalization.

Name that percussive tune: Associative memory and amplitude envelope.

A series of experiments demonstrated novel effects of amplitude envelope on associative memory, with tones exhibiting naturally decaying amplitude envelopes (e.g., those made by two wine glasses clinking) better associated with target objects than amplitude-invariant tones. In Experiment 1 participants learned associations between household objects and 4-note tone sequences constructed of spectrally matched pure tones with either "flat" or "percussive" amplitude envelopes. Those hearing percussive tones correctly recalled significantly more sequence-object associations. Experiment 2 demonstrated that participants hearing percussive tones learned the associations more quickly. Experiment 3 used "reverse percussive" tones (percussive tones played backwards) to test whether differences in overall energy might account for this effect, finding they did not lead to the same level of performance as percussive tones. Experiment 4 varied the envelope at encoding and retrieval to determine which stage of the task was most affected by the envelope manipulation. Participants hearing percussive tones at both encoding and retrieval performed significantly better than the other three groups (i.e., flat at encoding/percussive at retrieval, etc.). We conclude that amplitude envelope plays an important role in learning and memory, a finding with relevance to psychological research on audition and associative memory, as well as practical relevance for improving human-computer interface design.

Action-effects enhance explicit sequential learning.

Different studies have shown that action-effect associations seem to enhance implicit learning of motor sequences. In a recent study (Haider et al., Conscious Cognit 26:145-161, 2014), we found indications that action-effect learning might play a special role in acquiring explicit knowledge within an implicit learning situation. The current study aims at directly manipulating the action-effect contingencies in a Serial Reaction Time Task and examining its impact on explicit sequence knowledge. For this purpose, we created a situation in which the participants' responses led to a melodic tone sequence. For one group, these effect tones were contingently bound to the sequential responses and immediately followed the key press; for the second group, the tones were delayed by 400ms. For a third group, the tones also followed the response immediately and resulted in the same melody but were not contingently bound to the responses. A fourth control group received no effect tones at all. Only the group that experienced contingent effect tones that directly followed the response showed an increase in explicit sequence knowledge. The results are discussed in terms of the multi-modal structure of action-effect associations and the ideomotor principle of action control.

Pre-target neural oscillations predict variability in the detection of small pitch changes.

Pitch discrimination is important for language or music processing. Previous studies indicate that auditory perception depends on pre-target neural activity. However, so far the pre-target electrophysiological conditions which enable the detection of small pitch changes are not well studied, but might yield important insights into pitch-processing. We used magnetoencephalography (MEG) source imaging to reveal the pre-target effects of successful auditory detection of small pitch deviations from a sequence of standard tones. Participants heard a sequence of four pure tones and had to determine whether the last target tone was different or identical to the first three standard sounds. We found that successful pitch change detection could be predicted from the amplitude of theta (4–8 Hz) oscillatory activity in the right inferior frontal gyrus (IFG) as well as beta (12–30 Hz) oscillatory activity in the right auditory cortex. These findings confirm and extend evidence for the involvement of theta as well as beta-band activity in auditory perception.

Neural correlates of distraction and conflict resolution for nonverbal auditory events

In everyday situations auditory selective attention requires listeners to suppress task-irrelevant stimuli and to resolve conflicting information in order to make appropriate goal-directed decisions. Traditionally, these two processes (i.e. distractor suppression and conflict resolution) have been studied separately. In the present study we measured neuroelectric activity while participants performed a new paradigm in which both processes are quantified. In separate block of trials, participants indicate whether two sequential tones share the same pitch or location depending on the block’s instruction. For the distraction measure, a positive component peaking at ~250 ms was found – a distraction positivity. Brain electrical source analysis of this component suggests different generators when listeners attended to frequency and location, with the distraction by location more posterior than the distraction by frequency, providing support for the dual-pathway theory. For the conflict resolution measure, a negative frontocentral component (270–450 ms) was found, which showed similarities with that of prior studies on auditory and visual conflict resolution tasks. The timing and distribution are consistent with two distinct neural processes with suppression of task-irrelevant information occurring before conflict resolution. This new paradigm may prove useful in clinical populations to assess impairments in filtering out task-irrelevant information and/or resolving conflicting information.

A Dynamical Model of Pitch Memory Provides an Improved Basis for Implied Harmony Estimation.

Tonal melody can imply vertical harmony through a sequence of tones. Current methods for automatic chord estimation commonly use chroma-based features extracted from audio signals. However, the implied harmony of unaccompanied melodies can be difficult to estimate on the basis of chroma content in the presence of frequent nonchord tones. Here we present a novel approach to automatic chord estimation based on the human perception of pitch sequences. We use cohesion and inhibition between pitches in auditory short-term memory to differentiate chord tones and nonchord tones in tonal melodies. We model short-term pitch memory as a gradient frequency neural network, which is a biologically realistic model of auditory neural processing. The model is a dynamical system consisting of a network of tonotopically tuned nonlinear oscillators driven by audio signals. The oscillators interact with each other through nonlinear resonance and lateral inhibition, and the pattern of oscillatory traces emerging from the interactions is taken as a measure of pitch salience. We test the model with a collection of unaccompanied tonal melodies to evaluate it as a feature extractor for chord estimation. We show that chord tones are selectively enhanced in the response of the model, thereby increasing the accuracy of implied harmony estimation. We also find that, like other existing features for chord estimation, the performance of the model can be improved by using segmented input signals. We discuss possible ways to expand the present model into a full chord estimation system within the dynamical systems framework.

Artificial grammar learning in tamarins (Saguinus oedipus) in varying stimulus contexts.

The human ability to detect regularities in sound sequences is a fundamental substrate of our language faculty. However, is this an ability exclusive to human language processing, or have we usurped a more general learning mechanism for this purpose, one shared with other species? The current study is an attempt to replicate and extend Hauser, Weiss, and Marcus's (2002) retracted study (2010) of artificial grammar learning in tamarins to determine if tamarins can detect an underlying grammatical structure in a pattern of sounds. Human language consonant-vowel (CV) combinations from Hauser et al.'s original study, newly created tone sequences, and newly created monkey vocalizations made into sequences were used to familiarize tamarins to an AAB or ABB pattern. Tests of novel sounds in each condition were presented that either were consistent with the familiarized pattern or were different from it. Longer looking times toward the sound source (an audio speaker with a specific location in the auditory field) indicated recognition of novelty. Tamarins looked toward the speaker significantly longer with inconsistent human language CV sequences and with inconsistent tone sequences but not when an inconsistent monkey vocalization was presented. Moreover, tamarins showed differential rates of habituation to the different types of sound patterns, with more robust habituation to CV sequences and tone sequences than to monkey call sequences. The implications of these findings for the generality of learning mechanisms for linguistic and nonlinguistic input across species and the importance of testing across various stimuli are discussed. (PsycINFO Database Record

Build-up effect of auditory streaming in budgerigars (Melopsittacus undulatus)

When listening to a rapid tone sequence of the form ABA-ABA-ABA-… (where A and B are two tones of different frequencies and “-” indicates a silence interval), listeners may either hear one coherent “gallop” of three tones grouped together or two separate auditory streams (one high frequency, one low frequency) appearing to come from two sound sources. Research on humans indicates that the tendency to perceive two streams can be built up as exposure time increases. Neural recordings in European starlings show build-up effects of auditory streaming too. A lack of behavioral data on the build-up effect in nonhumans make it difficult to draw parallels between animals and humans. The present research aims to behaviorally validate the build-up effect of auditory streaming and factors that may influence the effect in nonhuman animals. Four budgerigars were tested in a categorization task using operant conditioning. “Streaming” categorization increased as the frequency separation increased. Additionally, in some frequency separations, as the sequence duration increases, the probability of “streaming” categorizations reported by the budgerigars is higher. These results indicate that budgerigars experience a build-up effect of auditory streaming behaviorally, and this effect is influenced by, but may not be limited to, different frequency separations.

Effects of temporal coherence and signal-frequency uncertainty for tone detection in a random-frequency multi-tonal masker

Detection thresholds tend to be lower when the spectral and/or temporal characteristics of the signal are predictable than unpredictable, particularly in an unpredictable masker. The present study evaluated the detrimental effect of frequency uncertainty and the beneficial effect of temporal coherence for a pure-tone signal presented in a multi-tonal masker with unpredictable frequency sequences. Stimuli were composed of 80-ms tone bursts. The signal was 1, 2, 4, or 8 sequential tone bursts, and burst frequency was either fixed across trials or randomly varied across trials but fixed across bursts. The masker was composed of four streams of tone bursts; the frequency of each burst was randomly drawn from a uniform distribution 250-4000 Hz, with the caveat that synchronous masker and signal bursts were separated by 1/5 oct or more. Signal and masker bursts were synchronously gated, and the masker played continuously throughout a threshold estimation track. Thresholds tended to improve with increasing numbers of signal bursts and worsen with increasing signal frequency uncertainty. These factors appeared to interact; signal frequency uncertainty was less detrimental when the signal was composed of larger numbers of bursts. Preliminary models of detection will be discussed.

Frequency tagging to track the neural processing of contrast in fast, continuous sound sequences.

The human auditory system presents a remarkable ability to detect rapid changes in fast, continuous acoustic sequences, as best illustrated in speech and music. However, the neural processing of rapid auditory contrast remains largely unclear, probably due to the lack of methods to objectively dissociate the response components specifically related to the contrast from the other components in response to the sequence of fast continuous sounds. To overcome this issue, we tested a novel use of the frequency-tagging approach allowing contrast-specific neural responses to be tracked based on their expected frequencies. The EEG was recorded while participants listened to 40-s sequences of sounds presented at 8Hz. A tone or interaural time contrast was embedded every fifth sound (AAAAB), such that a response observed in the EEG at exactly 8 Hz/5 (1.6 Hz) or harmonics should be the signature of contrast processing by neural populations. Contrast-related responses were successfully identified, even in the case of very fine contrasts. Moreover, analysis of the time course of the responses revealed a stable amplitude over repetitions of the AAAAB patterns in the sequence, except for the response to perceptually salient contrasts that showed a buildup and decay across repetitions of the sounds. Overall, this new combination of frequency-tagging with an oddball design provides a valuable complement to the classic, transient, evoked potentials approach, especially in the context of rapid auditory information. Specifically, we provide objective evidence on the neural processing of contrast embedded in fast, continuous sound sequences.NEW & NOTEWORTHY Recent theories suggest that the basis of neurodevelopmental auditory disorders such as dyslexia might be an impaired processing of fast auditory changes, highlighting how the encoding of rapid acoustic information is critical for auditory communication. Here, we present a novel electrophysiological approach to capture in humans neural markers of contrasts in fast continuous tone sequences. Contrast-specific responses were successfully identified, even for very fine contrasts, providing direct insight on the encoding of rapid auditory information.

Revisiting the "enigma" of musicians with dyslexia: Auditory sequencing and speech abilities.

Previous research has suggested a link between musical training and auditory processing skills. Musicians have shown enhanced perception of auditory features critical to both music and speech, suggesting that this link extends beyond basic auditory processing. It remains unclear to what extent musicians who also have dyslexia show these specialized abilities, considering often-observed persistent deficits that coincide with reading impairments. The present study evaluated auditory sequencing and speech discrimination in 52 adults comprised of musicians with dyslexia, nonmusicians with dyslexia, and typical musicians. An auditory sequencing task measuring perceptual acuity for tone sequences of increasing length was administered. Furthermore, subjects were asked to discriminate synthesized syllable continua varying in acoustic components of speech necessary for intraphonemic discrimination, which included spectral (formant frequency) and temporal (voice onset time [VOT] and amplitude envelope) features. Results indicate that musicians with dyslexia did not significantly differ from typical musicians and performed better than nonmusicians with dyslexia for auditory sequencing as well as discrimination of spectral and VOT cues within syllable continua. However, typical musicians demonstrated superior performance relative to both groups with dyslexia for discrimination of syllables varying in amplitude information. These findings suggest a distinct profile of speech processing abilities in musicians with dyslexia, with specific weaknesses in discerning amplitude cues within speech. Because these difficulties seem to remain persistent in adults with dyslexia despite musical training, this study only partly supports the potential for musical training to enhance the auditory processing skills known to be crucial for literacy in individuals with dyslexia. (PsycINFO Database Record

An auditory illusion reveals the role of streaming in the temporal misallocation of perceptual objects.

This study investigates the neural correlates and processes underlying the ambiguous percept produced by a stimulus similar to Deutsch's 'octave illusion', in which each ear is presented with a sequence of alternating pure tones of low and high frequencies. The same sequence is presented to each ear, but in opposite phase, such that the left and right ears receive a high-low-high … and a low-high-low … pattern, respectively. Listeners generally report hearing the illusion of an alternating pattern of low and high tones, with all the low tones lateralized to one side and all the high tones lateralized to the other side. The current explanation of the illusion is that it reflects an illusory feature conjunction of pitch and perceived location. Using psychophysics and electroencephalogram measures, we test this and an alternative hypothesis involving synchronous and sequential stream segregation, and investigate potential neural correlates of the illusion. We find that the illusion of alternating tones arises from the synchronous tone pairs across ears rather than sequential tones in one ear, suggesting that the illusion involves a misattribution of time across perceptual streams, rather than a misattribution of location within a stream. The results provide new insights into the mechanisms of binaural streaming and synchronous sound segregation.This article is part of the themed issue 'Auditory and visual scene analysis'.

Intentional switching of auditory attention between long and short sequential tone patterns.

The current study focuses on auditory task switching, more precisely on switching attention between different temporal patterns of the same auditory stimulus. Tone sequences consisting of nine different pitch tones were presented aurally. Three repetitive short 3-tone patterns (local focus) were combined to a long pattern (global focus), and each could be either rising or falling, resulting in congruent or incongruent combinations. Participants were informed by a cue if they had to attend to the short or to the long pattern, and they indicated if the target pattern was rising or falling by pressing one of two keys. In two experiments, we investigated cued switches between the two attentional foci. Switch costs in reaction times and errors were observed when switching from the long to the short pattern but not when switching from the short to the long pattern. These asymmetric switch costs were reduced when participants had more time to prepare for the switch in a condition with a prolonged cue-stimulus interval. In addition, participants made more errors when global and local patterns did not correspond to each other (i.e., in incongruent trials) when attending to either of the patterns, but this congruency effect was not modulated by preparation time. The data suggest that the mechanisms of task goal prioritizing, as indicated by the asymmetric attention switch costs, are dissociable from those underlying stimulus selection, as indicated by the congruency effects.

Impaired auditory-to-motor entrainment in Parkinson’s disease

Several electrophysiological studies suggest that Parkinson's disease (PD) patients have a reduced tendency to entrain to regular environmental patterns. Here we investigate whether this reduced entrainment concerns a generalized deficit or is confined to movement-related activity, leaving sensory entrainment intact. Magnetoencephalography was recorded during a rhythmic auditory target detection task in 14 PD patients and 14 control subjects. Participants were instructed to press a button when hearing a target tone amid an isochronous sequence of standard tones. The variable pitch of standard tones indicated the probability of the next tone to be a target. In addition, targets were occasionally omitted to evaluate entrainment uncontaminated by stimulus effects. Response times were not significantly different between groups and both groups benefited equally from the predictive value of standard tones. Analyses of oscillatory beta power over auditory cortices showed equal entrainment to the tones in both groups. By contrast, oscillatory beta power and event-related fields demonstrated a reduced engagement of motor cortical areas in PD patients, expressed in the modulation depth of beta power, in the response to omitted stimuli, and in an absent motor area P300 effect. Together, these results show equally strong entrainment of neural activity over sensory areas in controls and patients, but, in patients, a deficient translation of the adjustment to the task rhythm to motor circuits. We suggest that the reduced activation reflects not merely altered resonance to rhythmic external events, but a compromised recruitment of an endogenous response reflecting internal rhythm generation.NEW & NOTEWORTHY Previous studies suggest that motor cortical activity in PD patients has a reduced tendency to entrain to regular environmental patterns. This study demonstrates that the deficient entrainment in PD concerns the motor system only, by showing equally strong entrainment of neural activity over sensory areas in controls and patients but, in patients, a deficient translation of this adjustment to the task rhythm to motor circuits.

Using binocular rivalry to tag foreground sounds: Towards an objective visual measure for auditory multistability

In binocular rivalry, paradigms have been proposed for unobtrusive moment-by-moment readout of observers' perceptual experience ("no-report paradigms"). Here, we take a first step to extend this concept to auditory multistability. Observers continuously reported which of two concurrent tone sequences they perceived in the foreground: high-pitch (1008 Hz) or low-pitch (400 Hz) tones. Interstimulus intervals were either fixed per sequence (Experiments 1 and 2) or random with tones alternating (Experiment 3). A horizontally drifting grating was presented to each eye; to induce binocular rivalry, gratings had distinct colors and motion directions. To associate each grating with one tone sequence, a pattern on the grating jumped vertically whenever the respective tone occurred. We found that the direction of the optokinetic nystagmus (OKN)-induced by the visually dominant grating-could be used to decode the tone (high/low) that was perceived in the foreground well above chance. This OKN-based readout improved after observers had gained experience with the auditory task (Experiments 1 and 2) and for simpler auditory tasks (Experiment 3). We found no evidence that the visual stimulus affected auditory multistability. Although decoding performance is still far from perfect, our paradigm may eventually provide a continuous estimate of the currently dominant percept in auditory multistability.

Stimulus Phase Locking of Cortical Oscillations for Rhythmic Tone Sequences in Rats

Humans can rapidly detect regular patterns (i.e., within few cycles) without any special attention to the acoustic environment. This suggests that human sensory systems are equipped with a powerful mechanism for automatically predicting forthcoming stimuli to detect regularity. It has recently been hypothesized that the neural basis of sensory predictions exists for not only what happens (predictive coding) but also when a particular stimulus occurs (predictive timing). Here, we hypothesize that the phases of neural oscillations are critical in predictive timing, and these oscillations are modulated in a band-specific manner when acoustic patterns become predictable, i.e., regular. A high-density microelectrode array (10 × 10 within 4 × 4 mm2) was used to characterize spatial patterns of band-specific oscillations when a random-tone sequence was switched to a regular-tone sequence. Increasing the regularity of the tone sequence enhanced phase locking in a band-specific manner, notwithstanding the type of the regular sound pattern. Gamma-band phase locking increased immediately after the transition from random to regular sequences, while beta-band phase locking gradually evolved with time after the transition. The amplitude of the tone-evoked response, in contrast, increased with frequency separation with respect to the prior tone, suggesting that the evoked-response amplitude encodes sequence information on a local scale, i.e., the local order of tones. The phase locking modulation spread widely over the auditory cortex, while the amplitude modulation was confined around the activation foci. Thus, our data suggest that oscillatory phase plays a more important role than amplitude in the neuronal detection of tone sequence regularity, which is closely related to predictive timing. Furthermore, band-specific contributions may support recent theories that gamma oscillations encode bottom-up prediction errors, whereas beta oscillations are involved in top-down prediction.

Information-Theoretic Properties of Auditory Sequences Dynamically Influence Expectation and Memory.

A basic function of cognition is to detect regularities in sensory input to facilitate the prediction and recognition of future events. It has been proposed that these implicit expectations arise from an internal predictive coding model, based on knowledge acquired through processes such as statistical learning, but it is unclear how different types of statistical information affect listeners' memory for auditory stimuli. We used a combination of behavioral and computational methods to investigate memory for non-linguistic auditory sequences. Participants repeatedly heard tone sequences varying systematically in their information-theoretic properties. Expectedness ratings of tones were collected during three listening sessions, and a recognition memory test was given after each session. Information-theoretic measures of sequential predictability significantly influenced listeners' expectedness ratings, and variations in these properties had a significant impact on memory performance. Predictable sequences yielded increasingly better memory performance with increasing exposure. Computational simulations using a probabilistic model of auditory expectation suggest that listeners dynamically formed a new, and increasingly accurate, implicit cognitive model of the information-theoretic structure of the sequences throughout the experimental session.

The predictability of a partner’s actions modulates the sense of joint agency

When people coordinate their actions with others, they experience a sense of joint agency, i.e., shared control over actions and their consequences. The current study examined whether the predictability of others’ actions modulates joint agency. Each participant coordinated with two confederate partners to produce tone sequences that matched a metronome pace. The timing of the confederates’ actions was manipulated so that one partner’s actions were highly predictable in time and the other’s less predictable. After each sequence, participants rated their experience of joint agency on a scale from shared to independent control. People felt more shared control when they coordinated with the more predictable partner, even after controlling for their own performance accuracy and variability. Thus, people rely on predictions of others’ actions to derive a sense of joint agency during interpersonal coordination.

Temporal coherence structure rapidly shapes neuronal interactions

Perception of segregated sources is essential in navigating cluttered acoustic environments. A basic mechanism to implement this process is the temporal coherence principle. It postulates that a signal is perceived as emitted from a single source only when all of its features are temporally modulated coherently, causing them to bind perceptually. Here we report on neural correlates of this process as rapidly reshaped interactions in primary auditory cortex, measured in three different ways: as changes in response rates, as adaptations of spectrotemporal receptive fields following stimulation by temporally coherent and incoherent tone sequences, and as changes in spiking correlations during the tone sequences. Responses, sensitivity and presumed connectivity were rapidly enhanced by synchronous stimuli, and suppressed by alternating (asynchronous) sounds, but only when the animals engaged in task performance and were attentive to the stimuli. Temporal coherence and attention are therefore both important factors in auditory scene analysis.

Subjective perceptual organization of a complex auditory scene

Empirical research on the sequential decomposition of an auditory scene primarily relies on interleaved sound mixtures of only two tone sequences (e.g., ABAB…). This oversimplifies the sound decomposition problem by limiting the number of putative perceptual organizations. The current study used a sound mixture composed of three different tones (ABCABC…) that could be perceptually organized in many different ways. Participants listened to these sequences and reported their subjective perception by continuously choosing one out of 12 visually presented perceptual organization alternatives. Different levels of frequency and spatial separation were implemented to check whether participants' perceptual reports would be systematic and plausible. As hypothesized, while perception switched back and forth in each condition between various perceptual alternatives (multistability), spatial as well as frequency separation generally raised the proportion of segregated and reduced the proportion of integrated alternatives. During segregated percepts, in contrast to the hypothesis, many participants had a tendency to perceive two streams in the foreground, rather than reporting alternatives with a clear foreground-background differentiation. Finally, participants perceived the organization with intermediate feature values (e.g., middle tones of the pattern) segregated in the foreground slightly less often than similar alternatives with outer feature values (e.g., higher tones).