The perception of octave pitch affinity and harmonic fusion have a common origin.

Musicians say that the pitches of tones with a frequency ratio of 2:1 (one octave) have a distinctive affinity, even if the tones do not have common spectral components. It has been suggested, however, that this affinity judgment has no biological basis and originates instead from an acculturation process ‒ the learning of musical rules unrelated to auditory physiology. We measured, in young amateur musicians, the perceptual detectability of octave mistunings for tones presented alternately (melodic condition) or simultaneously (harmonic condition). In the melodic condition, mistuning was detectable only by means of explicit pitch comparisons. In the harmonic condition, listeners could use a different and more efficient perceptual cue: in the absence of mistuning, the tones fused into a single sound percept; mistunings decreased fusion. Performance was globally better in the harmonic condition, in line with the hypothesis that listeners used a fusion cue in this condition; this hypothesis was also supported by results showing that an illusory simultaneity of the tones was much less advantageous than a real simultaneity. In the two conditions, mistuning detection was generally better for octave compressions than for octave stretchings. This asymmetry varied across listeners, but crucially the listener-specific asymmetries observed in the two conditions were highly correlated. Thus, the perception of the melodic octave appeared to be closely linked to the phenomenon of harmonic fusion. As harmonic fusion is thought to be determined by biological factors rather than factors related to musical culture or training, we argue that octave pitch affinity also has, at least in part, a biological basis.

Automatic Frequency-Shift Detection in the Auditory System: A Review of Psychophysical Findings.

The human brain has the task of binding successive sounds produced by the same acoustic source into a coherent perceptual stream, and binding must be selective when several sources are concurrently active. Binding appears to obey a principle of spectral proximity: pure tones close in frequency are more likely to be bound than pure tones with remote frequencies. It has been hypothesized that the binding process is realized by automatic "frequency-shift detectors" (FSDs), comparable to the detectors of spatial motion in the visual system. In 2005, this hypothesis was supported by a psychophysical study showing that human listeners are able to identify the direction of a frequency shift between two successive pure tones while the first of these tones is not audible individually due to informational masking by other tones presented synchronously. A number of variants of this study have been performed since 2005, in order to confirm the existence of FSDs, to characterize their properties, and to clarify as far as possible their neural underpinnings. The results obtained up to now suggest that the working of the FSDs exploits an implicit sensory memory which is powerful with respect to both capacity and retention time. Tones within chords can be perceptually enhanced by small frequency shifts, in a manner suggesting that the FSDs can serve in auditory scene analysis not only as binding tools but also, to a limited extent, as segregation tools.

Detecting temporal changes in acoustic scenes: The variable benefit of selective attention.

Four experiments investigated change detection in acoustic scenes consisting of a sum of five amplitude-modulated pure tones. As the tones were about 0.7 octave apart and were amplitude-modulated with different frequencies (in the range 2-32 Hz), they were perceived as separate streams. Listeners had to detect a change in the frequency (experiments 1 and 2) or the shape (experiments 3 and 4) of the modulation of one of the five tones, in the presence of an informative cue orienting selective attention either before the scene (pre-cue) or after it (post-cue). The changes left intensity unchanged and were not detectable in the spectral (tonotopic) domain. Performance was much better with pre-cues than with post-cues. Thus, change deafness was manifest in the absence of an appropriate focusing of attention when the change occurred, even though the streams and the changes to be detected were acoustically very simple (in contrast to the conditions used in previous demonstrations of change deafness). In one case, the results were consistent with a model based on the assumption that change detection was possible if and only if attention was endogenously focused on a single tone. However, it was also found that changes resulting in a steepening of amplitude rises were to some extent able to draw attention exogenously. Change detection was not markedly facilitated when the change produced a discontinuity in the modulation domain, contrary to what could be expected from the perspective of predictive coding.

The Effect of Cochlear Damage on the Sensitivity to Harmonicity.

OBJECTIVES: A sum of simultaneous pure tones with harmonic relationships (i.e., simple frequency ratios) is normally heard as a single sound, with a single pitch, even when its components are fully resolved in the auditory periphery. This perceptual phenomenon called "harmonic fusion" is thought to play an important role in auditory scene analysis as listeners often have to segregate simultaneous harmonic sounds with different fundamental frequencies. The present study explored the consequences of mild or moderate cochlear hearing loss for the sensitivity to harmonicity and the detection of inharmonicity. DESIGN: The subjects were 12 normal-hearing (NH) listeners and 8 listeners with cochlear hearing loss amounting to 30 to 50 dB (mean: 42 dB) from 0.25 to 3 kHz. In each subject, thresholds for the detection of a change in the frequency ratio of simultaneous pure tones were measured with an adaptive forced-choice procedure. The standard frequency ratio was either harmonic (2:1, i.e., one octave) or inharmonic (0.8 or 1.2 octaves). The tones were presented at a low sensation level (at most 15 dB) within broadband noise, to minimize their cochlear interactions. In the main experimental conditions, the frequency register of the stimuli varied randomly within each trial, so that subjects were forced to process frequency ratios to achieve good performance; frequency discrimination was not sufficient. In other conditions, by contrast, frequency discrimination was sufficient to perform the task optimally. RESULTS: For both groups of subjects, thresholds in the main experimental conditions were lower (i.e., better) when the standard frequency ratio was harmonic than when it was inharmonic. This effect, revealing sensitivity to harmonicity, was weak for some members of the hearing-impaired group, but could be observed even in subjects showing a very poor frequency discrimination ability. The two groups, however, differed from each other with respect to the detection of inharmonicity: for the NH group, in agreement with previous results, negative deviations from one octave (i.e., compressions of this frequency ratio) were better detected than positive deviations (stretchings); for the hearing-impaired group, on the other hand, the sign of the deviations had no effect on performance. CONCLUSIONS: Sensitivity to harmonicity appears to be remarkably robust. However, it can be reduced in some listeners with mild or moderate cochlear damage. Moreover, as inharmonicity detection is asymmetric for NH listeners but apparently becomes symmetric in case of cochlear damage, it may be that listeners with cochlear damage do not detect inharmonicity in the same manner as NH listeners do. In some circumstances, inharmonicity can be detected on the basis of "beat" cues available in single frequency channels; however, the subjects tested here were unlikely to use cues of this type.

Pitch priming in sequences of two sounds.

Frequency discrimination limens (FDLs) were measured for pairs of stimuli differing from each other with respect to pitch salience. One of the two stimuli to be compared within a trial was a pure tone of at least 100 ms, evoking a salient pitch, while the other stimulus consisted of only eight sinusoidal cycles (experiment 1), or was a noise band with a Gaussian spectral envelope, evoking a weak pitch corresponding to the peak frequency (experiment 2). From trial to trial, frequency was varied randomly and widely. In both experiments, the FDLs were lower, by an average factor of about 3, when the stimulus with the more salient pitch preceded the other stimulus than vice versa. Evidence is presented against an interpretation of this temporal asymmetry in terms of memory limitations. It is suggested that the asymmetry reflects a pitch-priming effect. In two additional experiments, both of the stimuli to be compared within a trial were very short tone bursts or noise bands; perceptually, they differed only with respect to pitch height. Performance was markedly better than in experiments 1 and 2, and was not improved when the two stimuli were preceded by a 300-ms tone intended to produce pitch priming.

Harmonic fusion and pitch affinity: Is there a direct link?

Simultaneous pure tones approximately one octave apart tend to be fused perceptually and to evoke a single pitch sensation. Besides, sequentially presented pure tones show a subjective "affinity" or similarity in pitch when their frequency ratio is close to one octave. The aim of the study reported here was to determine if these two perceptual phenomena are directly related. Each stimulus was a triplet of simultaneous or successive pure tones forming frequency ratios varying across stimuli between 0.96 and 1.04 octaves. The tones were presented at a low sensation level (15 dB) within broadband threshold-equalizing noise, in order to prevent them from interacting in the cochlea when they were simultaneous. A large set of stimulus comparisons made by 18 listeners indicated that: (1) when the tones were simultaneous, maximal fusion was obtained for a mean frequency ratio deviating by less than 0.2% from one octave, and fusion decreased less rapidly above this frequency ratio than below it; (2) when the tones were presented successively, maximal pitch affinity was obtained for a mean frequency ratio significantly larger than one octave, and pitch affinity decreased more rapidly above this frequency ratio than below it. The differences between the results obtained for simultaneous and successive tones suggest that harmonic fusion and pitch affinity are unrelated phenomena.

Auditory attention is divisible: segregated tone streams can be tracked simultaneously.

Can auditory attention be split? We addressed this question using rapid sequences of tones alternating in frequency between 2 remote registers. In these rapid sequences, consecutive tones could not be perceptually linked; the tones were instead inevitably segregated into 2 concurrent melodic streams. Listeners had to determine if the 2 melodies interleaved in a sequence were exact transpositions of each other or not. This task could be performed successfully. More crucially, performance was better when each component tone of 1 melody was immediately transposed in the other melody than when component i of 1 melody was a transposition of component i-1 of the other melody. Nevertheless, because the melodies were segregated, listeners were unable to determine which was the leading melody when 2 interleaved melodies were immediate transpositions of each other. Our results are inconsistent with the hypothesis that listeners compared concurrent melodic streams using a memory-based serial-processing strategy. It instead appears that listeners were able to track such streams in parallel. Therefore, attention can be split between concurrent sensory streams even when the physical entities making up these streams do not overlap in time.

A late-emerging auditory deficit in autism.

OBJECTIVE: Individuals with autism spectrum disorders (ASD) show enhanced perceptual and memory abilities in the domain of pitch, but also perceptual deficits in other auditory domains. The present study investigated their skills with respect to "echoic memory," a form of short-term sensory memory intimately tied to auditory perception, using a developmental perspective. METHOD: We tested 23 high-functioning participants with ASD and 26 typically developing (TD) participants, distributed in two age groups (children vs. young adults; mean ages: ∼11 and ∼21 years). By means of an adaptive psychophysical procedure, we measured the longest period for which periodic (i.e., repeated) noise could be reliably discriminated from nonperiodic (i.e., plain random) noise. On each experimental trial, a single noise sample was presented to the participant, who had to classify this sound as periodic or nonperiodic. RESULTS: The TD adults performed, on average, much better than the other three groups, who performed similarly overall. As a function of practice, the measured thresholds improved for the TD participants, but did not change for the ASD participants. Thresholds were not correlated to performance in a test assessing verbal memory. The variance of the participants' response biases was larger among the ASD participants than among the TD participants. CONCLUSION: The results mainly suggest that echoic memory takes a long time to fully develop in TD humans, and that this development stops prematurely in persons with ASD.

The auditory enhancement effect is not reflected in the 80-Hz auditory steady-state response.

The perceptual salience of a target tone presented in a multitone background is increased by the presentation of a precursor sound consisting of the multitone background alone. It has been proposed that this "enhancement" phenomenon results from an effective amplification of the neural response to the target tone. In this study, we tested this hypothesis in humans, by comparing the auditory steady-state response (ASSR) to a target tone that was enhanced by a precursor sound with the ASSR to a target tone that was not enhanced. In order to record neural responses originating in the brainstem, the ASSR was elicited by amplitude modulating the target tone at a frequency close to 80 Hz. The results did not show evidence of an amplified neural response to enhanced tones. In a control condition, we measured the ASSR to a target tone that, instead of being perceptually enhanced by a precursor sound, was acoustically increased in level. This level increase matched the magnitude of enhancement estimated psychophysically with a forward masking paradigm in a previous experimental phase. We found that the ASSR to the tone acoustically increased in level was significantly greater than the ASSR to the tone enhanced by the precursor sound. Overall, our results suggest that the enhancement effect cannot be explained by an amplified neural response at the level of the brainstem. However, an alternative possibility is that brainstem neurons with enhanced responses do not contribute to the scalp-recorded ASSR.

No Need for Templates in the Auditory Enhancement Effect.

The audibility of a target tone in a multitone background masker is enhanced by the presentation of a precursor sound consisting of the masker alone. There is evidence that precursor-induced neural adaptation plays a role in this perceptual enhancement. However, the precursor may also be strategically used by listeners as a spectral template of the following masker to better segregate it from the target. In the present study, we tested this hypothesis by measuring the audibility of a target tone in a multitone masker after the presentation of precursors which, in some conditions, were made dissimilar to the masker by gating their components asynchronously. The precursor and the following sound were presented either to the same ear or to opposite ears. In either case, we found no significant difference in the amount of enhancement produced by synchronous and asynchronous precursors. In a second experiment, listeners had to judge whether a synchronous multitone complex contained exactly the same tones as a preceding precursor complex or had one tone less. In this experiment, listeners performed significantly better with synchronous than with asynchronous precursors, showing that asynchronous precursors were poorer perceptual templates of the synchronous multitone complexes. Overall, our findings indicate that precursor-induced auditory enhancement cannot be fully explained by the strategic use of the precursor as a template of the following masker. Our results are consistent with an explanation of enhancement based on selective neural adaptation taking place at a central locus of the auditory system.

Enhancement of increments in spectral amplitude: further evidence for a mechanism based on central adaptation.

The threshold for detecting a tone in a multitone masker is lower when the masker-plus-signal stimulus is preceded by a copy of the masker. One potential explanation of this "enhancement" phenomenon is that the -precursor stimulus acts as a "template" of the subsequent masker, thus helping listeners to segregate the signal from the masker. To assess this idea, we measured enhancement for precursors that were perceptually similar to the masker and for precursors that were made dissimilar to the masker by gating their components asynchronously. We found that the two types of precursor produced similar amounts of enhancement. This was true not only when the precursor and the subsequent test stimulus were presented to the same ear but also when they were presented to opposite ears. In a second experiment, we checked that the precursors with asynchronously gated components were perceptually poor templates of the subsequent maskers. Listeners now had to discriminate between test stimuli -containing the same components as the precursor and test stimuli containing all but one of the precursor components. We found that in this experimental situation, where enhancement could play no role, gating the precursor components asynchronously disrupted performance. Overall, our results are inconsistent with the hypothesis that precursors producing enhancement are beneficial because they are used as perceptual templates of the masker. Our results are instead consistent with an -explanation of enhancement based on selective neural adaptation taking place at a central locus of the auditory system.

The perceptual enhancement of tones by frequency shifts.

In a chord of pure tones with a flat spectral profile, one tone can be perceptually enhanced relative to the other tones by the previous presentation of a slightly different chord. "Intensity enhancement" (IE) is obtained when the component tones of the two chords have the same frequencies, but in the first chord the target of enhancement is attenuated relative to the other tones. "Frequency enhancement" (FE) is obtained when both chords have a flat spectral profile, but the target of enhancement shifts in frequency from the first to the second chord. We report here an experiment in which IE and FE were measured using a task requiring the listener to indicate whether or not the second chord included a tone identical to a subsequent probe tone. The results showed that a global attenuation of the first chord relative to the second chord disrupted IE more than FE. This suggests that the mechanisms of IE and FE are not the same. In accordance with this suggestion, computations of the auditory excitation patterns produced by the chords indicate that the mechanism of IE is not sufficient to explain FE for small frequency shifts.

Auditory discrimination of frequency ratios: the octave singularity.

Sensitivity to frequency ratios is essential for the perceptual processing of complex sounds and the appreciation of music. This study assessed the effect of ratio simplicity on ratio discrimination for pure tones presented either simultaneously or sequentially. Each stimulus consisted of four 100-ms pure tones, equally spaced in terms of frequency ratio and presented at a low intensity to limit interactions in the auditory periphery. Listeners had to discriminate between a reference frequency ratio of 0.97 octave (about 1.96:1) and target frequency ratios, which were larger than the reference. In the simultaneous condition, the obtained psychometric functions were nonmonotonic: as the target frequency ratio increased from 0.98 octave to 1.04 octaves, discrimination performance initially increased, then decreased, and then increased again; performance was better when the target was exactly one octave (2:1) than when the target was slightly larger. In the sequential condition, by contrast, the psychometric functions were monotonic and there was no effect of frequency ratio simplicity. A control experiment verified that the non-monotonicity observed in the simultaneous condition did not originate from peripheral interactions between the tones. Our results indicate that simultaneous octaves are recognized as "special" frequency intervals by a mechanism that is insensitive to the sign (positive or negative) of deviations from the octave, whereas this is apparently not the case for sequential octaves.

Auditory enhancement of increments in spectral amplitude stems from more than one source.

A component of a test sound consisting of simultaneous pure tones perceptually "pops out" if the test sound is preceded by a copy of itself with that component attenuated. Although this "enhancement" effect was initially thought to be purely monaural, it is also observable when the test sound and the precursor sound are presented contralaterally (i.e., to opposite ears). In experiment 1, we assessed the magnitude of ipsilateral and contralateral enhancement as a function of the time interval between the precursor and test sounds (10, 100, or 600 ms). The test sound, randomly transposed in frequency from trial to trial, was followed by a probe tone, either matched or mismatched in frequency to the test sound component which was the target of enhancement. Listeners' ability to discriminate matched probes from mismatched probes was taken as an index of enhancement magnitude. The results showed that enhancement decays more rapidly for ipsilateral than for contralateral precursors, suggesting that ipsilateral enhancement and contralateral enhancement stem from at least partly different sources. It could be hypothesized that, in experiment 1, contralateral precursors were effective only because they provided attentional cues about the target tone frequency. In experiment 2, this hypothesis was tested by presenting the probe tone before the precursor sound rather than after the test sound. Although the probe tone was then serving as a frequency cue, contralateral precursors were again found to produce enhancement. This indicates that contralateral enhancement cannot be explained by cuing alone and is a genuine sensory phenomenon.

A note about insensitivity to pitch-change direction.

Some listeners are insensitive to the direction of pure-tone frequency changes when the standard frequency is roved widely over trials, but less so when the standard frequency is fixed and trial-by-trial feedback is provided. The present experiment tested the hypothesis that fixing the standard frequency and providing feedback is advantageous for direction-impaired listeners because under these conditions the listeners can learn to respond correctly without genuinely perceiving frequency-change direction. This hypothesis was ruled out by the experiment. It appears instead that direction-impaired listeners find it difficult to ignore the irrelevant frequency changes introduced by roving.

Frequency-shift detectors bind binaural as well as monaural frequency representations.

Previous psychophysical work provided evidence for the existence of automatic frequency-shift detectors (FSDs) that establish perceptual links between successive sounds. In this study, we investigated the characteristics of the FSDs with respect to the binaural system. Listeners were presented with sound sequences consisting of a chord of pure tones followed by a single test tone. Two tasks were performed. In the "present/absent" task, the test tone was either identical to one of the chord components or positioned halfway in frequency between two components, and listeners had to discriminate between these two possibilities. In the "up/down" task, the test tone was slightly different in frequency from one of the chord components and listeners had to identify the direction (up or down) of the corresponding shift. When the test tone was a pure tone presented monaurally, either to the same ear as the chord or to the opposite ear, listeners performed the up/down task better than the present/absent task. This paradoxical advantage for directional frequency shifts, providing evidence for FSDs, persisted when the test tone was replaced by a dichotic stimulus consisting of noise but evoking a pitch sensation as a consequence of binaural processing. Performance in the up/down task was similar for the dichotic stimulus and for a monaural narrow-band noise matched in pitch salience to it. Our results indicate that the FSDs are insensitive to sound localization mechanisms and operate on central frequency representations, at or above the level of convergence of the monaural auditory pathways.

Enhancing a tone by shifting its frequency or intensity.

When a test sound consisting of pure tones with equal intensities is preceded by a precursor sound identical to the test sound except for a reduction in the intensity of one tone, an auditory "enhancement" phenomenon occurs: In the test sound, the tone which was previously softer stands out perceptually. Here, enhancement was investigated using inharmonic sounds made up of five pure tones well resolved in the auditory periphery. It was found that enhancement can be elicited not only by increases in intensity but also by shifts in frequency. In both cases, when the precursor and test sounds are separated by a 500-ms delay, inserting a burst of pink noise during the delay has little effect on enhancement. Presenting the precursor and test sounds to opposite ears rather than to the same ear significantly reduces the enhancement resulting from increases in intensity, but not the enhancement resulting from shifts in frequency. This difference suggests that the mechanisms of enhancement are not identical for the two types of change. For frequency shifts, enhancement may be partly based on the existence of automatic "frequency-shift detectors" [Demany and Ramos, J. Acoust. Soc. Am. 117, 833-841 (2005)].

Implicit versus explicit frequency comparisons: two mechanisms of auditory change detection.

Listeners had to compare, with respect to pitch (frequency), a pure tone (T) to a combination of pure tones presented subsequently (C). The elements of C were either synchronous, and therefore difficult to hear out individually, or asynchronous and therefore easier to hear out individually. In the "present/absent" condition, listeners had to judge if T reappeared in C or not. In the "up/down" condition, the task was to judge if the element of C most similar to T was higher or lower than T. When the elements of C were synchronous, the up/down task was found to be easier than the present/absent task; the converse result was obtained when the elements of C were asynchronous. This provides evidence for a duality of auditory comparisons between tone frequencies: (1) implicit comparisons made by automatic and direction-sensitive "frequency-shift detectors"; (2) explicit comparisons more sensitive to the magnitude of a frequency change than to its direction. Another experiment suggests that although the frequency-shift detectors cannot compare effectively two tones separated by an interfering tone, they are largely insensitive to interfering noise bursts.

Fundamental differences in change detection between vision and audition.

We compared auditory change detection to visual change detection using closely matched stimuli and tasks in the two modalities. On each trial, participants were presented with a test stimulus consisting of ten elements: pure tones with various frequencies for audition, or dots with various spatial positions for vision. The test stimulus was preceded or followed by a probe stimulus consisting of a single element, and two change-detection tasks were performed. In the "present/absent" task, the probe either matched one randomly selected element of the test stimulus or none of them; participants reported present or absent. In the "direction-judgment" task, the probe was always slightly shifted relative to one randomly selected element of the test stimulus; participants reported the direction of the shift. Whereas visual performance was systematically better in the present/absent task than in the direction-judgment task, the opposite was true for auditory performance. Moreover, whereas visual performance was strongly dependent on selective attention and on the time interval separating the probe from the test stimulus, this was not the case for auditory performance. Our results show that small auditory changes can be detected automatically across relatively long temporal gaps, using an implicit memory system that seems to have no similar counterpart in the visual domain.

Tuning properties of the auditory frequency-shift detectors.

Demany and Ramos [(2005). J. Acoust. Soc. Am. 117, 833-841] found that it is possible to hear an upward or downward pitch change between two successive pure tones differing in frequency even when the first tone is informationally masked by other tones, preventing a conscious perception of its pitch. This provides evidence for the existence of automatic frequency-shift detectors (FSDs) in the auditory system. The present study was intended to estimate the magnitude of the frequency shifts optimally detected by the FSDs. Listeners were presented with sound sequences consisting of (1) a 300-ms or 100-ms random "chord" of synchronous pure tones, separated by constant intervals of either 650 cents or 1000 cents; (2) an interstimulus interval (ISI) varying from 100 to 900 ms; (3) a single pure tone at a variable frequency distance (Delta) from a randomly selected component of the chord. The task was to indicate if the final pure tone was higher or lower than the nearest component of the chord. Irrespective of the chord's properties and of the ISI, performance was best when Delta was equal to about 120 cents (1/10 octave). Therefore, this interval seems to be the frequency shift optimally detected by the FSDs.