What's special about human speech? A student exercise for comparing speech production between humans and chimpanzees.

Modern humans and chimpanzees share a common ancestor on the phylogenetic tree, yet chimpanzees do not spontaneously produce speech or speech sounds. The lab exercise presented in this paper was developed for undergraduate students in a course entitled "What's Special About Human Speech?" The exercise is based on acoustic analyses of the words "cup" and "papa" as spoken by Viki, a home-raised, speech-trained chimpanzee, as well as the words spoken by a human. The analyses allow students to relate differences in articulation and vocal abilities between Viki and humans to the known anatomical differences in their vocal systems. Anatomical and articulation differences between humans and Viki include (1) potential tongue movements, (2) presence or absence of laryngeal air sacs, (3) presence or absence of vocal membranes, and (4) exhalation vs inhalation during production.

Cochlear tuning and the peripheral representation of harmonic sounds in mammals.

Albert Feng was a prominent comparative neurophysiologist whose research provided numerous contributions towards understanding how the spectral and temporal characteristics of vocalizations underlie sound communication in frogs and bats. The present study is dedicated to Al's memory and compares the spectral and temporal representations of stochastic, complex sounds which underlie the perception of pitch strength in humans and chinchillas. Specifically, the pitch strengths of these stochastic sounds differ between humans and chinchillas, suggesting that humans and chinchillas may be using different cues. Outputs of auditory filterbank models based on human and chinchilla cochlear tuning were examined. Excitation patterns of harmonics are enhanced in humans as compared with chinchillas. In contrast, summary correlograms are degraded in humans as compared with chinchillas. Comparing summary correlograms and excitation patterns with corresponding behavioral data on pitch strength suggests that the dominant cue for pitch strength in humans is spectral (i.e., harmonic) structure, whereas the dominant cue for chinchillas is temporal (i.e., envelope) structure. The results support arguments that the broader cochlear tuning in non-human mammals emphasizes temporal cues for pitch perception, whereas the sharper cochlear tuning in humans emphasizes spectral cues.

Perception of degraded speech by chinchillas (Chinchilla laniger): Word-level stimulus generalization.

One characteristic of human speech perception is a remarkable ability to recognize speech when the speech signal is highly degraded. It has been argued that this ability to perceive highly degraded speech reflects speech-specific mechanisms. The present study tested this hypothesis by measuring the ability of chinchillas to recognize noise-vocoded (NV) versions of naturally spoken monosyllabic words using operant conditioning in a stimulus generalization paradigm. Chinchillas do not generalize the vocoded words to be perceptually equivalent to the naturally spoken words. The responses from chinchillas to the vocoded words fall well below their responses to the naturally spoken words. In this case, pitch cues rather than speech cues may be controlling the behavioral responses. To reduce pitch cues, chinchillas were retrained using 64-channel NV words. The responses from chinchillas to the vocoded test words were now similar to those of the 64-channel versions and were similar to those obtained from human listeners. However, responses obtained from chinchillas to time-reversed versions were high and similar to responses obtained to time-normal versions suggesting that the cue controlling behavioral responses was the phonetic structure of the words. These results show that chinchillas used different acoustic cues than human listeners. The ability of chinchillas to recognize NV words as being perceptually equivalent to the naturally spoken versions is inferior compared to that of human listeners. The findings suggest that the ability of human listeners to recognize highly degraded speech is unlikely to be based solely on the general auditory and perceptual mechanisms that are common among mammals. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Perception of noise-vocoded tone complexes: A time domain analysis based on an auditory filterbank model.

When a wideband harmonic tone complex (wHTC) is passed through a noise vocoder, the resulting sounds can have spectra with large peak-to-valley ratios, but little or no periodicity strength in the autocorrelation functions. We measured judgments of pitch strength for normal-hearing listeners for noise-vocoded wideband harmonic tone complexes (NV-wHTCs) relative to standard and anchor stimuli. The standard was a 1-channel NV-wHTC and the anchor was either the unprocessed wHTC or an infinitely-iterated rippled noise (IIRN). Although there is variability among individuals, the magnitude judgment functions obtained with the IIRN anchor suggest different listening strategies among individuals. In order to gain some insight into possible listening strategies, test stimuli were analyzed at the output of an auditory filterbank model based on gammatone filters. The weak periodicity strengths of NV-wHTCs observed in the stimulus autocorrelation functions are augmented at the output of the gammatone filterbank model. Six analytical models of pitch strength were evaluated based on summary correlograms obtained from the gammatone tone filterbank. The results of the filterbank analysis suggest that, contrary to the weak or absent periodicity strengths in the stimulus domain, temporal cues contribute to pitch strength perception of noise-vocoded harmonic stimuli such that listeners' judgments of pitch strength reflect a nonlinear, weighted average of the temporal information between the fine structure and the envelope.

Acoustic analysis of the frequency-dependent coupling between the frog's ears.

The ears of anurans are coupled through the Eustachian tubes and mouth cavity. The degree of coupling varies with frequency showing a bandpass characteristic, but the characteristics differ between empirically measured data based on auditory nerve responses and tympanic membrane vibration. In the present study, the coupling was modeled acoustically as a tube connected with a side branch. This tube corresponds to the Eustachian tubes, whereas the side branch corresponds to the mouth cavity and nares. The analysis accounts for the frequency dependency shown by the empirical data and reconciles the differences observed between the coupling as measured by tympanic membrane vibration and auditory nerve responses.

Perception of degraded speech sounds differs in chinchilla and human listeners.

The behavioral responses of chinchillas to noise-vocoded versions of naturally spoken speech sounds were measured using stimulus generalization and operant conditioning. Behavioral performance for speech generalization by chinchillas is compared to recognition by a group of human listeners for the identical speech sounds. The ability of chinchillas to generalize the vocoded versions as tokens of the natural speech sounds is far less than recognition by human listeners. In many cases, responses of chinchillas to noise-vocoded speech sounds were more similar to responses to band limited noise than to the responses to natural speech sounds. Chinchillas were also tested with a middle C musical note as played on a piano. Comparison of the responses of chinchillas for the middle C condition to the responses obtained for the speech conditions suggest that chinchillas may be more influenced by fundamental frequency than by formant structure. The differences between vocoded speech perception in chinchillas and human listeners may reflect differences in their abilities to resolve the formants along the cochlea. It is argued that lengthening of the cochlea during human evolution may have provided one of the auditory mechanisms that influenced the evolution of speech-specific mechanisms.

Processing pitch in a nonhuman mammal (Chinchilla laniger).

Whether the mechanisms giving rise to pitch reflect spectral or temporal processing has long been debated. Generally, sounds having strong harmonic structures in their spectra have strong periodicities in their temporal structures. We found that when a wideband harmonic tone complex is passed through a noise vocoder, the resulting sound can have a harmonic structure with a large peak-to-valley ratio, but with little or no periodicity in the temporal structure. To test the role of harmonic structure in pitch perception for a nonhuman mammal, we measured behavioral responses to noise-vocoded tone complexes in chinchillas (Chinchilla laniger) using a stimulus generalization paradigm. Chinchillas discriminated either a harmonic tone complex or an iterated rippled noise from a 1-channel vocoded version of the tone complex. When tested with vocoded versions generated with 8, 16, 32, 64, and 128 channels, responses were similar to those of the 1-channel version. Behavioral responses could not be accounted for based on harmonic peak-to-valley ratio as the acoustic cue, but could be accounted for based on temporal properties of the autocorrelation functions such as periodicity strength or the height of the first peak. The results suggest that pitch perception does not arise through spectral processing in nonhuman mammals but rather through temporal processing. The conclusion that spectral processing contributes little to pitch in nonhuman mammals may reflect broader cochlear tuning than that described in humans.

Pitch strength of noise-vocoded harmonic tone complexes in normal-hearing listeners.

To study the role of harmonic structure in pitch perception, normal-hearing listeners were tested using noise-vocoded harmonic tone complexes. When tested in a magnitude judgment procedure using vocoded versions generated with 2-128 channels, judgments of pitch strength increased systematically as the number of channels increased and reflected acoustic cues based on harmonic peak-to-valley ratio, but not cues based on periodicity strength. When tested in a fundamental frequency discrimination task, listeners correctly recognized the direction of pitch change with as few as eight noise-vocoded channels. The results suggest that spectral processing contributes substantially to pitch perception in normal-hearing listeners.

Perception of the missing fundamental by chinchillas in the presence of low-pass masking noise.

The pitch of the missing fundamental (F0) is one of the principal psychological attributes of human pitch perception. Behavioral responses to harmonic tone complexes having missing F0s were measured in chinchillas using operant conditioning and stimulus generalization. Animals were trained to discriminate between tone complexes having a 500-Hz F0 and a 125-Hz F0. When animals were tested with tone complexes having the same F0s, but where the F0s were missing, responses were similar to those obtained when the F0s were present, suggesting that missing F0 sounds were perceptually equivalent to F0 present sounds. Behavioral responses to F0 present and missing F0 stimuli were similar in the presence of low-pass masking noise, suggesting that the perception was not due to the reinsertion of the F0 through cochlear nonlinearities. Gradients in behavioral responses were observed when the F0s of test complexes were systematically varied, suggesting the existence of a psychological dimension related to F0. Behavioral responses were related to the F0 rather than to spectral differences among test stimuli when the F0 and spectrum were varied independently. The results indicate that chinchillas possess a pitch-like perception of the missing F0 that is unlikely to arise from cochlear distortion products.

Critical bands and critical ratios in animal psychoacoustics: an example using chinchilla data.

This paper suggests that critical ratios obtained in noise-masked tone studies are not good indicators of critical bandwidths obtained in both human and nonhuman animal subjects. A probe-tone detection study using chinchilla subjects suggests that they may be broadband processors in detection tasks as opposed to human subjects who use narrow-band, critical-band processing. If chinchilla and other nonhuman animal subjects are wideband processors, this can partially explain why their critical ratios are significantly greater than those measured in human subjects. Thus, large critical ratios obtained for nonhuman animals may indicate processing inefficiency rather than wide critical bands.

Representation of the spectral dominance region of pitch in the steady-state temporal discharge patterns of cochlear nucleus units.

Single-unit responses to infinitely iterated rippled noise and wideband noise were recorded from the cochlear nucleus of anesthetized chinchillas. Rippled noises had a fixed delay of 4 ms, and spectral depth was varied by attenuating the delayed version of the noise. Temporal discharge patterns were analyzed using neural autocorrelograms, and responses to rippled noises were compared to wideband noise responses. Chopper units with best frequencies in the range of the first to second harmonics of the rippled noise showed large differences in discharge patterns between rippled noise and wideband noise responses, but chopper units with best frequencies centered at higher harmonics did not show large differences. Consequently, the Chopper group of units showed no evidence of a neural representation of the dominance region. Primarylike units did show a neural representation of dominance that is related to behavioral performance. For Primarylike units with best frequencies around the third to fifth harmonics of the rippled noise, large differences in discharge patterns between rippled noise and wideband noise responses were observed. The results suggest that bushy cells play an important role in processing pitch-related information and should be included as important elements in neural models of periodicity processing.

Pitch perception in chinchillas (Chinchilla laniger): stimulus generalization using rippled noise.

Rippled noises evoke the perception of pitch in human listeners. Infinitely iterated rippled noise (IIRN) is generated when wideband noise (WBN) is delayed, attenuated, and added to the original WBN through either a positive (+) or a negative (-) feedback loop. The pitch of IIRN[+] is matched to the reciprocal of the delay, whereas the pitch of IIRN[-] for the same delay is an octave lower. Chinchillas (Chinchilla laniger) were trained to discriminate IIRN[+] with a 4-ms delay from IIRN[+] with a 2-ms delay and then tested in a stimulus generalization paradigm with IIRN[+] at delays between 2 and 4 ms. Systematic gradients in behavioral response occurred along the dimension of delay, suggesting that a perceptual dimension corresponding to pitch exists for IIRN[+]. Behavioral responses to IIRN[-] test stimuli were more variable among chinchillas, suggesting that IIRN[-] did not evoke similar pitches relative to IIRN[+]. Systematic gradients in behavioral response were observed when IIRN[-] test stimuli were presented in the context of other IIRN[-] stimuli. Thus, other perceptual cues such as timbre may dominate the pitch cues when IIRN[-] test stimuli are presented in the context of IIRN[+] stimuli.

Pitch cue learning in chinchillas: the role of spectral region in the training stimulus.

Chinchillas were trained to discriminate a cosine-phase harmonic tone complex (COS) from wideband noise (WBN) and tested in a stimulus generalization paradigm with tone complexes in which phase differed between frequency regions. In this split-phase condition, responses to complexes made of random-phase low frequencies, cosine-phase high frequencies were similar to responses to the COS-training stimulus. However, responses to complexes made of cosine-phase low frequencies, random-phase high frequencies were generally lower than their responses to the COS-training stimulus. When tested with sine-phase (SIN) and random-phase (RND) tone complexes, responses were large for SIN, but were small for RND. Chinchillas were then trained to discriminate infinitely-iterated rippled noise (IIRN) from WBN and tested with noises in which the spectral ripple differed between frequency regions. In this split-spectrum condition, responses were large to noises made of rippled-spectrum low frequencies, flat-spectrum high frequencies, whereas responses were generally lower to noises made of flat-spectrum low frequencies, rippled-spectrum high frequencies. The results suggest that chinchillas listen across all frequencies, but attend to high frequencies when discriminating COS from WBN and attend to low frequencies when discriminating IIRN from WBN.

Listening experience with iterated rippled noise alters the perception of 'pitch' strength of complex sounds in the chinchilla.

Behavioral responses obtained from chinchillas trained to discriminate a cosine-phase harmonic tone complex from wideband noise indicate that the perception of 'pitch' strength in chinchillas is largely influenced by periodicity information in the stimulus envelope. The perception of 'pitch' strength was examined in chinchillas in a stimulus generalization paradigm after animals had been retrained to discriminate infinitely iterated rippled noise from wideband noise. Retrained chinchillas gave larger behavioral responses to test stimuli having strong fine structure periodicity, but weak envelope periodicity. That is, chinchillas learn to use the information in the fine structure and consequently, their perception of 'pitch' strength is altered. Behavioral responses to rippled noises having similar periodicity strengths, but large spectral differences were also tested. Responses to these rippled noises were similar, suggesting a temporal analysis can be used to account for the behavior. Animals were then retested using the cosine-phase harmonic tone complex as the expected signal stimulus. Generalization gradients returned to those obtained originally in the naïve condition, suggesting that chinchillas do not remain "fine structure listeners," but rather revert back to being "envelope listeners" when the periodicity strength in the envelope of the expected stimulus is high.

Perception of the periodicity strength of complex sounds by the chinchilla.

The perception of periodicity strength was studied in chinchillas using a stimulus generalization paradigm in an operant-conditioning, positive reinforcement behavioral task. Stimuli consisted of cosine-phase and random-phase harmonic complex tones, infinitely iterated rippled noises, and wideband noise. These stimuli vary in periodicity strength as measured by autocorrelation functions and are known to generate a continuum in the perception of pitch strength in human listeners. Chinchillas were trained to discriminate a cosine-phase harmonic tone complex from wideband noise and tested in the generalization paradigm using random-phase tone complexes and iterated rippled noises as probe stimuli. Chinchillas were tested in three different conditions in which the periods of the fundamental frequencies of the tone complexes were fixed at 2 ms, 4 ms, or 8 ms. Behavioral responses obtained from chinchillas were related to stimulus periodicity strength. For most animals, the behavioral responses to random-phase tone complexes were smaller than those to cosine-phase tone complexes. The behavioral responses were analyzed in terms of the Auditory Image Model of Patterson et al. [Patterson, R.D., Allerhand, M.H., Giguère, C., J. Acoust. Soc. Am. 98 (1995) 1890-1894], and the results suggest that the periodicity information in the stimulus envelope has a large influence in controlling the behavioral response of the chinchilla. Comparison of the generalization data obtained in the present study to magnitude estimation data obtained previously in human subjects suggests a greater influence of stimulus envelope for the perception of periodicity strength in chinchillas than for the perception of pitch strength in human listeners.

Pitch strength and Stevens's power law.

Recent studies have suggested that the saliency or the strength of pitch of complex sounds can be accounted for on the basis of the temporal properties in the stimulus waveform as measured by the height of the first peak in the waveform autocorrelation function. We used a scaling procedure to measure the pitch strength from 15 listeners for four different pitches of complex sounds in which the height of the first peak in the autocorrelation function systematically varied. Pitch strength judgments were evaluated in terms of a modification of Stevens's power law in which temporal information was used from both the waveform fine structure and the envelope. Best fits of this modified power law to the judged pitch strengths indicate that the exponent in Stevens's power law is greater than 1. The results suggest that pitch strength is primarily determined by the waveform fine structure, but the stimulus envelope can also contribute to the pitch strength.