3MRC Cognition and Brain Sciences Unit

Acoustic sequences such as speech and music are generally perceived as coherent auditory “streams,” which can be individually attended to and followed over time. Although the psychophysical stimulus parameters governing this “auditory streaming ” are well established, the brain mechanisms underlying the formation of auditory streams remain largely unknown. In particular, an essential feature of the phenomenon, which corresponds to the fact that the segregation of sounds into streams typically takes several seconds to build up, remains unexplained. Here, we show that this and other major features of auditorystream formation measured in humans using alternating-tone sequences can be quantitatively accounted for based on single-unit responses recorded in the primary auditory cortex (A1) of awake rhesus monkeys listening to the same sound sequences

Neural Correlates of Auditory Perceptual Awareness under Informational Masking

Our ability to detect target sounds in complex acoustic backgrounds is often limited not by the ear's resolution, but by the brain's information-processing capacity. The neural mechanisms and loci of this “informational masking” are unknown. We combined magnetoencephalography with simultaneous behavioral measures in humans to investigate neural correlates of informational masking and auditory perceptual awareness in the auditory cortex. Cortical responses were sorted according to whether or not target sounds were detected by the listener in a complex, randomly varying multi-tone background known to produce informational masking. Detected target sounds elicited a prominent, long-latency response (50–250 ms), whereas undetected targets did not. In contrast, both detected and undetected targets produced equally robust auditory middle-latency, steady-state responses, presumably from the primary auditory cortex. These findings indicate that neural correlates of auditory awareness in informational masking emerge between early and late stages of processing within the auditory cortex

Auditory Stream Segregation and the Perception of Across-Frequency Synchrony

This study explored the extent to which sequential auditory grouping affects the perception of temporal synchrony. In Experiment 1, listeners discriminated between 2 pairs of asynchronous "target" tones at different frequencies, A and B, in which the B tone either led or lagged. Thresholds were markedly higher when the target tones were temporally surrounded by "captor tones" at the A frequency than when the captor tones were absent or at a remote frequency. Experiment 2 extended these findings to asynchrony detection, revealing that the perception of synchrony, one of the most potent cues for simultaneous auditory grouping, is not immune to competing effects of sequential grouping. Experiment 3 examined the influence of ear separation on the interactions between sequential and simultaneous grouping cues. The results showed that, although ear separation could facilitate perceptual segregation and impair asynchrony detection, it did not prevent the perceptual integration of simultaneous sounds. Keywords: perceptual organization, auditory perception, stream segregation, asynchrony Sensitivity to seemingly low-level sensory features in a scene can be profoundly influenced by the way in which the scene is perceptually organized. A striking example of this in visual perception relates to the finding by In the auditory modality, one of the most compelling examples of the influence of perceptual organization on sensitivity comes from demonstrations that listeners are largely unable to correctly perceive the relative timing of sounds that form part of separate perceptual "streams." The first lines of evidence for this were provided b

Relative Pitch Perception and the Detection of Deviant Tone Patterns.

Most people are able to recognise familiar tunes even when played in a different key. It is assumed that this depends on a general capacity for relative pitch perception; the ability to recognise the pattern of inter-note intervals that characterises the tune. However, when healthy adults are required to detect rare deviant melodic patterns in a sequence of randomly transposed standard patterns they perform close to chance. Musically experienced participants perform better than naïve participants, but even they find the task difficult, despite the fact that musical education includes training in interval recognition.To understand the source of this difficulty we designed an experiment to explore the relative influence of the size of within-pattern intervals and between-pattern transpositions on detecting deviant melodic patterns. We found that task difficulty increases when patterns contain large intervals (5-7 semitones) rather than small intervals (1-3 semitones). While task difficulty increases substantially when transpositions are introduced, the effect of transposition size (large vs small) is weaker. Increasing the range of permissible intervals to be used also makes the task more difficult. Furthermore, providing an initial exact repetition followed by subsequent transpositions does not improve performance. Although musical training correlates with task performance, we find no evidence that violations to musical intervals important in Western music (i.e. the perfect fifth or fourth) are more easily detected. In summary, relative pitch perception does not appear to be conducive to simple explanations based exclusively on invariant physical ratios

Signal detection in animal psychoacoustics: analysis and simulation of sensory and decision-related influences

Signal detection theory (SDT) provides a framework for interpreting psychophysical experiments, separating the putative internal sensory representation and the decision process. SDT was used to analyse ferret behavioural responses in a (yes–no) tone-in-noise detection task. Instead of measuring the receiver-operating characteristic (ROC), we tested SDT by comparing responses collected using two common psychophysical data collection methods. These (Constant Stimuli, Limits) differ in the set of signal levels presented within and across behavioural sessions. The results support the use of SDT as a method of analysis: SDT sensory component was unchanged between the two methods, even though decisions depended on the stimuli presented within a behavioural session. Decision criterion varied trial-by-trial: a ‘yes’ response was more likely after a correct rejection trial than a hit trial. Simulation using an SDT model with several decision components reproduced the experimental observations accurately, leaving only ∼10% of the variance unaccounted for. The model also showed that trial-by-trial dependencies were unlikely to influence measured psychometric functions or thresholds. An additional model component suggested that inattention did not contribute substantially. Further analysis showed that ferrets were changing their decision criteria, almost optimally, to maximise the reward obtained in a session. The data suggest trial-by-trial reward-driven optimization of the decision process. Understanding the factors determining behavioural responses is important for correlating neural activity and behaviour. SDT provides a good account of animal psychoacoustics, and can be validated using standard psychophysical methods and computer simulations, without recourse to ROC measurements

Dimension-specific attention directs learning and listening on auditory training tasks

The relative contributions of bottom-up versus top-down sensory inputs to auditory learning are not well established. In our experiment, listeners were instructed to perform either a frequency discrimination (FD) task ("FD-train group") or an intensity discrimination (ID) task ("ID-train group") during training on a set of physically identical tones that were impossible to discriminate consistently above chance, allowing us to vary top-down attention whilst keeping bottom-up inputs fixed. A third, control group did not receive any training. Only the FD-train group improved on a FD probe following training, whereas all groups improved on ID following training. However, only the ID-train group also showed changes in performance accuracy as a function of interval with training on the ID task. These findings suggest that top-down, dimension-specific attention can direct auditory learning, even when this learning is not reflected in conventional performance measures of threshold change

Informational masking and the effects of differences in fundamental frequency and fundamental-frequency contour on phonetic integration in a formant ensemble

This study explored the effects on speech intelligibility of across-formant differences in fundamental frequency (ΔF0) and F0 contour. Sentence-length speech analogues were presented dichotically (left=F1+F3; right=F2), either alone or—because competition usually reveals grouping cues most clearly—accompanied in the left ear by a competitor for F2 (F2C) that listeners must reject to optimize recognition. F2C was created by inverting the F2 frequency contour. In experiment 1, all left-ear formants shared the same constant F0 and ΔF0F2 was 0 or ±4 semitones. In experiment 2, all left-ear formants shared the natural F0 contour and that for F2 was natural, constant, exaggerated, or inverted. Adding F2C lowered keyword scores, presumably because of informational masking. The results for experiment 1 were complicated by effects associated with the direction of ΔF0F2; this problem was avoided in experiment 2 because all four F0 contours had the same geometric mean frequency. When the target formants were presented alone, scores were relatively high and did not depend on the F0F2 contour. F2C impact was greater when F2 had a different F0 contour from the other formants. This effect was a direct consequence of the associated ΔF0; the F0F2 contour per se did not influence competitor impact

The contribution of visual information to the perception of speech in noise with and without informative temporal fine structure

Understanding what is said in demanding listening situations is assisted greatly by looking at the face of a talker. Previous studies have observed that normal-hearing listeners can benefit from this visual information when a talker’s voice is presented in background noise. These benefits have also been observed in quiet listening conditions in cochlear-implant users, whose device does not convey the informative temporal fine structure cues in speech, and when normal-hearing individuals listen to speech processed to remove these informative temporal fine structure cues. The current study (1) characterised the benefits of visual information when listening in background noise; and (2) used sine-wave vocoding to compare the size of the visual benefit when speech is presented with or without informative temporal fine structure. The accuracy with which normal-hearing individuals reported words in spoken sentences was assessed across three experiments. The availability of visual information and informative temporal fine structure cues was varied within and across the experiments. The results showed that visual benefit was observed using open- and closed-set tests of speech perception. The size of the benefit increased when informative temporal fine structure cues were removed. This finding suggests that visual information may play an important role in the ability of cochlear-implant users to understand speech in many everyday situations. Models of audio-visual integration were able to account for the additional benefit of visual information when speech was degraded and suggested that auditory and visual information was being integrated in a similar way in all conditions. The modelling results were consistent with the notion that audio-visual benefit is derived from the optimal combination of auditory and visual sensory cues

Effect of stimulus type and pitch salience on pitch-sequence processing

Using a same-different discrimination task, it has been shown that discrimination performance for sequences of complex tones varying just detectably in pitch is less dependent on sequence length (1, 2, or 4 elements) when the tones contain resolved harmonics than when they do not [Cousineau, Demany, and Pessnitzer (2009). J. Acoust. Soc. Am. 126, 3179-3187]. This effect had been attributed to the activation of automatic frequency-shift detectors (FSDs) by the shifts in resolved harmonics. The present study provides evidence against this hypothesis by showing that the sequence-processing advantage found for complex tones with resolved harmonics is not found for pure tones or other sounds supposed to activate FSDs (narrow bands of noise and wide-band noises eliciting pitch sensations due to interaural phase shifts). The present results also indicate that for pitch sequences, processing performance is largely unrelated to pitch salience per se: for a fixed level of discriminability between sequence elements, sequences of elements with salient pitches are not necessarily better processed than sequences of elements with less salient pitches. An ideal-observer model for the same-different binary-sequence discrimination task is also developed in the present study. The model allows the computation of d' for this task using numerical methods

Stream segregation in the anesthetized auditory cortex

Auditory stream segregation describes the way that sounds are perceptually segregated into groups or streams on the basis of perceptual attributes such as pitch or spectral content. For sequences of pure tones, segregation depends on the tones' proximity in frequency and time. In the auditory cortex (and elsewhere) responses to sequences of tones are dependent on stimulus conditions in a similar way to the perception of these stimuli. However, although highly dependent on stimulus conditions, perception is also clearly influenced by factors unrelated to the stimulus, such as attention. Exactly how ‘bottom-up’ sensory processes and non-sensory ‘top-down’ influences interact is still not clear. Here, we recorded responses to alternating tones (ABAB …) of varying frequency difference (FD) and rate of presentation (PR) in the auditory cortex of anesthetized guinea-pigs. These data complement previous studies, in that top-down processing resulting from conscious perception should be absent or at least considerably attenuated. Under anesthesia, the responses of cortical neurons to the tone sequences adapted rapidly, in a manner sensitive to both the FD and PR of the sequences. While the responses to tones at frequencies more distant from neuron best frequencies (BFs) decreased as the FD increased, the responses to tones near to BF increased, consistent with a release from adaptation, or forward suppression. Increases in PR resulted in reductions in responses to all tones, but the reduction was greater for tones further from BF. Although asymptotically adapted responses to tones showed behavior that was qualitatively consistent with perceptual stream segregation, responses reached asymptote within 2 s, and responses to all tones were very weak at high PRs (>12 tones per second). A signal-detection model, driven by the cortical population response, made decisions that were dependent on both FD and PR in ways consistent with perceptual stream segregation. This included showing a range of conditions over which decisions could be made either in favor of perceptual integration or segregation, depending on the model ‘decision criterion’. However, the rate of ‘build-up’ was more rapid than seen perceptually, and at high PR responses to tones were sometimes so weak as to be undetectable by the model. Under anesthesia, adaptation occurs rapidly, and at high PRs tones are generally poorly represented, which compromises the interpretation of the experiment. However, within these limitations, these results complement experiments in awake animals and humans. They generally support the hypothesis that ‘bottom-up’ sensory processing plays a major role in perceptual organization, and that processes underlying stream segregation are active in the absence of attention