Frequency and intensity discrimination in humans and monkeys

Frequency and intensity DLs were compared in humans and monkeys using a repeating standard "yes-no" procedure in which subjects reported frequency increments, frequency decrements, intensity increments, or intensity decrements in an ongoing train of 1.0-kHz tone bursts. There was only one experimental condition (intensity increments) in which monkey DLs (1.5-2.0 dB) overlapped those of humans (1.0-1.8 dB). For discrimination of both increments and decrements in frequency, monkey DLs (16-33 Hz) were approximately seven times larger than those of humans (2.4-4.8 Hz), and for discrimination of intensity decrements, monkey DLs (4.4-7.0 dB) were very unstable and larger than those of humans (1.0-1.8 dB). For intensity increment discrimination, humans and monkeys also exhibited similar DLs as SL was varied. However, for frequency increment discrimination, best DLs for humans occurred at a high (50 dB) SL, whereas best DLs for monkeys occurred at a moderate (30 dB) SL. Results are discussed in terms of various neural mechanisms that might be differentially engaged by humans and monkeys in performing these tasks; for example, different amounts of temporal versus rate coding in frequency discrimination, and different mechanisms for monitoring rate decreases in intensity discrimination. The implications of these data for using monkeys as models of human speech sound discrimination are also discussed.     

Discrimination of frequency transitions by human infants

Difference limens (DLs) for linear frequency transitions using a 1.0-kHz pulsed-tone standard were obtained from 6- to 9-month-old human infants in a series of three experiments. A repeating standard "yes-no" operant headturning technique and an adaptive staircase (tracking) procedure were used to obtain difference limens from a total of 71 infants. The DLs for 300-ms upward and downward linear frequency sweeps were approximately 3%-4% when the repeating standard was an unmodulated 1.0-kHz pulsed tone of 300-ms duration. These DLs for frequency sweeps were not significantly different from DLs for frequency increments and decrements using 330-ms pulsed tones [J. M. Sinnott and R. N. Aslin, J. Acoust. Soc. Am. 78, 1986-1992 (1985)]. The DLs for frequency sweeps of 50 ms appended to the beginning or the end of a 250-ms unmodulated 1.0-kHz tone were approximately 6%-7%. This greater DL for brief frequency sweeps was confirmed by varying the duration but not the extent of the sweep. Finally, DLs were greater than 50% when the repeating standard was a 50-ms rising or falling frequency sweep appended to the beginning of a 250-ms unmodulated 1.0-kHz tone. These results suggest that rapid frequency transitions are much more difficult to discriminate from frequency transitions of the same category (rising or falling) than from either a frequency transition of the opposite category (falling or rising) or an unmodulated tone.(ABSTRACT TRUNCATED AT 250 WORDS)     

Sound intensity processing by the goldfish

Capacities of the goldfish for intensity discrimination were studied using classical respiratory conditioning and a staircase psychophysical procedure. Physiological studies on single saccular (auditory) nerve fibers under similar stimulus conditions helped characterize the dimensions of neural activity used in intensity discrimination. Incremental intensity difference limens (IDLs in dB) for 160-ms increments in continuous noise, 500-ms noise bursts, and 500-ms, 800-Hz tone bursts are 2 to 3 dB, are independent of overall level, and vary with signal duration according to a power function with a slope averaging - 0.33. Noise decrements are relatively poorly detected and the silent gap detection threshold is about 35 ms. The IDLs for increments and decrements in an 800-Hz continuous tone are about 0.13 dB, are independent of duration, and are level dependent. Unlike mammalian auditory nerve fibers, some goldfish saccular fibers show variation in recovery time to tonal increments and decrements, and adaptation to a zero rate. Unit responses to tone increments and decrements show rate effects generally in accord with previous observations on intracellular epsp's in goldfish saccular fibers. Neurophysiological correlates of psychophysical intensity discrimination data suggest the following: (1) noise gap detection may be based on spike rate increments which follow gap offset; (2) detection of increments and decrements in continuous tones may be determined by steep low-pass filtering in peripheral neural channels which enhance the effects of spectral "splatter" toward the lower frequencies; (3) IDLs for pulsed signals of different duration can be predicted from the slopes of rate-intensity functions and spike rate variability in individual auditory nerve fibers; and (4) at different sound pressure levels, different populations of peripheral fibers provide the information used in intensity discrimination.     

Detection of time- and bandlimited increments and decrements in a random-level noise.

The purpose of this study was to compare detection of increments and decrements occurring over limited regions of time and frequency within a 500-ms broadband (0-6000 Hz) noise. Three listeners tracked detection thresholds adaptively in a two-interval, two-alternative forced-choice task. Thresholds were measured for both increments and decrements in level [delta L = 10 log10(1 + delta N0/N0) dB, where N0 is the spectral power density of the noise] as a function of signal duration (T = 30-500 ms) for a range of signal bandwidths (W = 62-6000 Hz) that were logarithmically centered around 2500 Hz. Listeners were forced to rely on temporal- and spectral-profile cues for detection due to randomization of overall presentation level from interval to interval, which rendered overall energy an inconsistent cue. Increments were detectable for all combinations of W and T, whereas decrements were not consistently detectable for W < 500 Hz. Narrow-band decrements were not detectable due to spread of excitation from the spectral edges of the noise into the decrements. Increment and decrement thresholds were similar for W > or = 1000 Hz. Temporal- and spectral-integration effects were observed for both increments and decrements. The exceptions were for random-level conditions in which the signal matched the bandwidth or duration of the standard. A multicue decision process is described qualitatively to explain how the combination of temporal- and spectral-profile cues can produce temporal- and spectral-integration effects in the absence of overall-energy cues.     

Auditory duration discrimination in Old World monkeys (Macaca, Cercopithecus) and humans

Auditory duration DLs at 2.0 kHz were measured in Old World monkeys (Macaca, Cercopithecus) and humans using a go, no-go repeating standard AX procedure and positive reinforcement operant conditioning techniques. For a 200-ms standard, monkey DLs were 45-125 ms, compared to 15-27 ms for humans. Weber fractions (delta T/T) for all species were smallest at standard durations of 200-400 ms and increased as standard duration decreased to 25 ms. Varying intensity from 30-70 dB SPL had only minor effects on DLs, except at the lowest levels tested, where DLs were elevated slightly. Monkeys had difficulty discriminating duration decrements, in contrast to humans. Results are discussed in relation to other comparative psychoacoustic data and primate vocal communication, including human speech.     

An autocorrelation model with place dependence to account for the effect of harmonic number on fundamental frequency discrimination

Fundamental frequency (f0) difference limens (DLs) were measured as a function of f0 for sine- and random-phase harmonic complexes, bandpass filtered with 3-dB cutoff frequencies of 2.5 and 3.5 kHz (low region) or 5 and 7 kHz (high region), and presented at an average 15 dB sensation level (approximately 48 dB SPL) per component in a wideband background noise. Fundamental frequencies ranged from 50 to 300 Hz and 100 to 600 Hz in the low and high spectral regions, respectively. In each spectral region, f0 DLs improved dramatically with increasing f0 as approximately the tenth harmonic appeared in the passband. Generally, f0 DLs for complexes with similar harmonic numbers were similar in the two spectral regions. The dependence of f0 discrimination on harmonic number presents a significant challenge to autocorrelation (AC) models of pitch, in which predictions generally depend more on spectral region than harmonic number. A modification involving a "lag window"is proposed and tested, restricting the AC representation to a limited range of lags relative to each channel's characteristic frequency. This modified unitary pitch model was able to account for the dependence of f0 DLs on harmonic number, although this correct behavior was not based on peripheral harmonic resolvability.     

Intensity discrimination, increment detection, and magnitude estimation for 1-kHz tones

Intensity difference limens (DLs) were measured over a wide intensity range for 200-ms, 1-kHz gated tones and for 200-ms increments in continuous 1-kHz tones. Magnitude estimates also were obtained for the gated tones over a comparable intensity range. The discrimination data are in general agreement with those from earlier studies but they extend them by showing: (1) good discrimination for gated tones over at least a 115-dB dynamic range; (2) a slight increase in the relative DL (delta I/I) as intensity increases above 95 dB SPL; (3) smaller DLs for increments than for gated tones, with the difference approximately independent of intensity; (4) negligible "negative masking" when thresholds are expressed as intensity differences (delta I). For two of the three subjects, magnitude estimates do not conform to a single-exponent power law for suprathreshold intensities. Over the middle range of intensities where a single exponent is appropriate, the value of the exponent is less than 0.1 for all subjects.     

Pure-tone sensitivity of human infants

Pure-tone thresholds at frequencies ranging from 250 to 8000 Hz were estimated for 3-, 6-, and 12-month-old infants and for adults, using the Observer-based Psychoacoustic Procedure (OPP). Sounds were presented monaurally using an earphone. Psychometric functions of infants were similar to those of adults, although 3-month-olds had shallower functions at higher frequencies. The thresholds of 6- and 12-month-old infants were 10-15 dB higher than those of the adults, with the difference being greater at lower frequencies. This result is in general agreement with results from other laboratories. The thresholds of 3-month-olds were 15-30 dB higher than those of adults. The greatest difference between 3-month-olds and adults was at 8000 Hz. This threshold difference is smaller than that reported in earlier behavioral studies; higher thresholds at high frequencies have been previously reported for newborn and 3-month-old infants. The relative contributions of sensory and nonsensory variables to these age differences are discussed.     

Siamang gibbons exceed the saccular threshold: intensity of the song of Hylobates syndactylus

Measurements are reported of the intensity of the siamang gibbon loud call obtained from the vocal bouts of three family groups at Twycross Zoo, UK. Across 25 samples the maximum intensity ranged from 95 to 113 dB SPL (linear frequency-weighting and fast time-weighting) and exhibited three frequency modes of 250-315 Hz, 630-800 Hz and 1.2-1.6 kHz. The lowest frequency mode, which may correspond to the "boom" sound produced by resonance of the siamang inflated vocal sac, had a mean maximum intensity of 99 dB SPL. These values, which are in excess of the saccular acoustic threshold of about 90 dB at 300 Hz for air conducted sound, suggest that primate loud calls recruit a primitive mode of acoustic sensitivity furnished by the sacculus. Thus reproductive vocal behavior of primates may be influenced by a primitive acoustical reward pathway inherited from a common ancestor with anamniotes. In humans such a pathway could explain the compulsion for exposure to loud music.     

Pitch perception of concurrent harmonic tones with overlapping spectra

Fundamental frequency difference limens (F0DLs) were measured for a target harmonic complex tone with nominal fundamental frequency (F0) of 200 Hz, in the presence and absence of a harmonic masker with overlapping spectrum. The F0 of the masker was 0, ± 3, or ± 6 semitones relative to 200 Hz. The stimuli were bandpass filtered into three regions: 0-1000 Hz (low, L), 1600-2400 Hz (medium, M), and 2800-3600 Hz (high, H), and a background noise was used to mask combination tones and to limit the audibility of components falling on the filter skirts. The components of the target or masker started either in cosine or random phase. Generally, the effect of F0 difference between target and masker was small. For the target alone, F0DLs were larger for random than cosine phase for region H. For the target plus masker, F0DLs were larger when the target had random phase than cosine phase for regions M and H. F0DLs increased with increasing center frequency of the bandpass filter. Modeling using excitation patterns and "summary autocorrelation" and "stabilized auditory image" models suggested that use of temporal fine structure information can account for the small F0DLs obtained when harmonics are barely, if at all, resolved.     

Differential sensitivity to vowel continua in Old World monkeys (Macaca) and humans

Previous studies indicate that monkey pure tone frequency discrimination is quantitatively and qualitatively very different from that of humans: Monkey DLs at 1.0 and 2.0 kHz are up to 20 times larger than human DLs, and monkeys DLs increase as sensation level increases, in contrast to human DLs [Sinnott et al., J. Acoust. Soc. Am. 78, 1977-1985 (1985); Sinnott et al., J. Comp. Psychol. 101, 126-131 (1987)]. These results led to an hypothesis that monkey frequency discrimination is more dependent upon "rate" coding than is that of humans. The present study compared monkey and human DLs for formant frequency changes along three synthetic vowel continua /I-i/, /ae-epsilon/, and /a-v/. Here, monkey DLs for formants near 1.0 and 2.0 kHz (32-48 Hz) were only about two to three times larger than human DLs (11-21 Hz), and both monkeys and humans exhibited relatively similar, flat sensation level functions. Taken together, these data indicate that monkey and human frequency discrimination is more similar in the case of a complex vowel stimulus than in the case of a simple pure tone stimulus. Results are discussed in relation to "rate" versus "temporal" coding of tones and vowels in the auditory system.     

Cortical, auditory, evoked potentials in response to changes of spectrum and amplitude

The acoustic change complex (ACC) is a scalp-recorded negative-positive voltage swing elicited by a change during an otherwise steady-state sound. The ACC was obtained from eight adults in response to changes of amplitude and/or spectral envelope at the temporal center of a three-formant synthetic vowel lasting 800 ms. In the absence of spectral change, the group mean waveforms showed a clear ACC to amplitude increments of 2 dB or more and decrements of 3 dB or more. In the presence of a change of second formant frequency (from perceived /u/ to perceived /i/), amplitude increments increased the magnitude of the ACC but amplitude decrements had little or no effect. The fact that the just detectable amplitude change is close to the psychoacoustic limits of the auditory system augurs well for the clinical application of the ACC. The failure to find a condition under which the spectrally elicited ACC is diminished by a small change of amplitude supports the conclusion that the observed ACC to a change of spectral envelope reflects some aspect of cortical frequency coding. Taken together, these findings support the potential value of the ACC as an objective index of auditory discrimination capacity.     

Level and age effects in infant frequency discrimination

Frequency difference limens (FDLs) were estimated for 3-, 6-, and 12-month-old infants and for adults using pure tones at 500, 1000, and 4000 Hz. Each listener provided an FDL at 40 dB and at a higher (80 dB, in most cases) sensation level (SL). An observer-based behavioral testing technique was used. The FDLs of 3-month-olds were worse than those of adults at all three frequencies, and increased with increasing frequency. The FDLs of 6- and 12-month-olds were worse than those of adults at 500 and 1000 Hz, but not at 4000 Hz. Decreasing the SL led to an increase in the FDL of about the same magnitude at all ages, and the same age differences were found at both SLs. Thus infant-adult differences in FDL are not a simple consequence of differences in absolute sensitivity. Infant FDLs at one SL were also found to be significantly correlated with the FDL at the other SL. The FDLs at one age were, in general, predictive of the FDL at a later age in a longitudinal sample of infants. Models that might account for these age-related differences are discussed.     

Effects of environmental noise on computer-derived voice estimates from female speakers

The purpose of this study was to examine the influence of noise on voice profile statistics from female samples. Six young adult females served as subjects. Five had normal voices; one had a pathological voice with accompanying bilateral vocal nodules. Each female subject was required to match a generated 235 Hz tone (+/- 2 Hz) while maintaining a constant output level of 70 dB SPL (+/- 5 dB). Data collected from a previous study involving a normal male subject were included for comparative purposes. Noise was generated from a personal computer fan which had a strong center frequency component at 235 Hz. Six different A-weighted signal-to-noise [S/N(A)] conditions were created, ranging in 5 dB increments from 25 to 0 dB. Results revealed that fundamental frequency was reasonably resistant to the effects of noise and to the effects of the noisy (pathological) voice signal. Jitter and shimmer estimates generally increased as noise floors elevated. The greatest amount of measurement error was found for the pathological female voice when captured in the presence of environmental noise. Findings are discussed relative to clinical issues surrounding measurement error.     

Frequency and intensity difference limens for harmonics within complex tones

A two-interval, two-alternative forced choice task was used to estimate frequency difference limens (DLs) for individual harmonics within complex tones, and DLs for the periodicity (i.e., number of periods per s) of the whole complexes. For complex tones with equal-amplitude harmonics, the DLs for the lowest harmonics were small (less than one percent). The DLs increased rather abruptly around the fifth to seventh harmonic. The highest harmonic in each complex was also well discriminated, and the discriminability of a single high harmonic was markedly improved by increasing its level relative to the other components. The DL for a complex tone was generally smaller than the frequency DL of its most discriminable component. The DL for a complex was found to be predictable from the DLs of the harmonics comprising the complex, using a formula derived by Goldstein [J. Acoust. Soc. Am. 54, 1496-1516 (1973)] from his optimum processor theory for the formation of the pitch of complex tones. The DL for a complex is sometimes primarily determined by high harmonics, such as the highest harmonic, or a harmonic whose level exceeds that of adjacent harmonics. We also measured intensity DLs for individual harmonics within complex tones. The intensity DLs were smallest for low harmonic numbers, and for the highest harmonic in a complex. An excitation-pattern model was used to determine whether the frequency DLs of harmonics within complex tones could be explained in terms of place mechanisms, i.e., in terms of changes in the amount of excitation at appropriate frequency places. We conclude that place mechanisms are not adequate, and that information about the frequencies of individual harmonics is probably carried in the time patterning of neural impulses.     

Effects of frequency and duration on psychometric functions for detection of increments and decrements in sinusoids in noise

Psychometric functions for detecting increments or decrements in level of sinusoidal pedestals were measured for increment and decrement durations of 5, 10, 20, 50, 100, and 200 ms and for frequencies of 250, 1000, and 4000 Hz. The sinusoids were presented in background noise intended to mask spectral splatter. A three-interval, three-alternative procedure was used. The results indicated that, for increments, the detectability index d' was approximately proportional to delta I/I. For decrements, d' was approximately proportional to delta L. The slopes of the psychometric functions increased (indicating better performance) with increasing frequency for both increments and decrements. For increments, the slopes increased with increasing increment duration up to 200 ms at 250 and 1000 Hz, but at 4000 Hz they increased only up to 50 ms. For decrements, the slopes increased for durations up to 50 ms, and then remained roughly constant, for all frequencies. For a center frequency of 250 Hz, the slopes of the psychometric functions for increment detection increased with duration more rapidly than predicted by a "multiple-looks" hypothesis, i.e., more rapidly than the square root of duration, for durations up to 50 ms. For center frequencies of 1000 and 4000 Hz, the slopes increased less rapidly than predicted by a multiple-looks hypothesis, for durations greater than about 20 ms. The slopes of the psychometric functions for decrement detection increased with decrement duration at a rate slightly greater than the square root of duration, for durations up to 50 ms, at all three frequencies. For greater durations, the increase in slope was less than proportional to the square root of duration. The results were analyzed using a model incorporating a simulated auditory filter, a compressive nonlinearity, a sliding temporal integrator, and a decision device based on a template mechanism. The model took into account the effects of both the external noise and an assumed internal noise. The model was able to account for the major features of the data for both increment and decrement detection.     

Acoustic response and tuning in saccular nerve fibers of the goldfish (Carassius auratus)

The acoustic frequency selectivity of over 500 saccular nerve fibers of the goldfish was studied using automated threshold tracking based on spike rate increments defined statistically. Saccular fibers of the goldfish show great variation in (1) best sensitivity (-26 to + 35 dB re: 1 dyn/cm2), (2) best frequency (below 100 to 1770 Hz), (3) spontaneous rate (0 to over 200 spikes/s), (4) spontaneous type (silent, regular, irregular, burst), and (5) degree of tuning (Q 10 dB from less than 0.1 to 2). Saccular fibers may be grouped into four nonoverlapping categories based on tuning and best frequency: (1) untuned (less than 10-dB variation in sensitivity between 100 and 1000 Hz), (2) low frequency (BF from below 120 to 290 Hz), (3) midfrequency (BF between 330 and 670 Hz), and (4) high frequency (BF between 790 and 1770 Hz). Within each category, all spontaneous rates and types, and all degrees of tuning can be observed. The least sensitive fibers within each group have zero spontaneous rates. The goldfish is like all other vertebrates studied in that the peripheral auditory system is adapted for frequency selectivity throughout the animal's entire frequency range of hearing. Peripheral tuning most likely accounts for behavioral determinations of the "auditory filter" and for the detectability of signals masked by noise. The signal-to-noise ratio enhancement provided by these peripheral filters is likely to be of primary biological significance. A "place principle" of sound quality analysis based on lines "labeled" according to best frequency in the brain cannot be ruled out on the basis of the peripheral physiology.     

Masking effects on ABR waves I and V in infants and adults

The effects of high- and low-pass masking on waves I and V of the auditory brain stem response (ABR) were measured in normal infants who were 2-4 weeks old, and in adults. The signal was a 4-kHz tone pip presented at 86 dB peak equivalent sound-pressure level (p.e.SPL). The masking patterns were different for latency and amplitude criteria, and were also different for infants and adults. The largest difference between infants and adults was seen in the wave I data. Low-pass maskers were very disruptive of the infant wave I, while little or no effect was noted on the adult wave I. High-pass maskers were very disruptive of the adult wave I, while less of an effect was measured on the infant wave I. The wave V data were similar between groups. Cochlear regions which contribute most importantly to wave I extend up to one octave above the frequency of the signal in adults, and to one-half octave above the signal frequency in infants. The reasons for the differences found between infants and adults are uncertain. Two possible mechanisms which can explain these data are differences in peripheral auditory sensitivity, and differences in tuning characteristics of the auditory system.     

The relationship between frequency selectivity and pitch discrimination: sensorineural hearing loss

This study tested the relationship between frequency selectivity and the minimum spacing between harmonics necessary for accurate fo discrimination. Fundamental frequency difference limens (fo DLs) were measured for ten listeners with moderate sensorineural hearing loss (SNHL) and three normal-hearing listeners for sine- and random-phase harmonic complexes, bandpass filtered between 1500 and 3500 Hz, with fo's ranging from 75 to 500 Hz (or higher). All listeners showed a transition between small (good) fo DLs at high fo's and large (poor) fo DLs at low fo's, although the fo at which this transition occurred (fo,tr) varied across listeners. Three measures thought to reflect frequency selectivity were significantly correlated to both the fo,tr and the minimum fo DL achieved at high fo's: (1) the maximum fo for which fo DLs were phase dependent, (2) the maximum modulation frequency for which amplitude modulation and quasi-frequency modulation were discriminable, and (3) the equivalent rectangular bandwidth of the auditory filter, estimated using the notched-noise method. These results provide evidence of a relationship between fo discrimination performance and frequency selectivity in listeners with SNHL, supporting "spectral" and "spectro-temporal" theories of pitch perception that rely on sharp tuning in the auditory periphery to accurately extract fo information.     

Discrimination of dynamic interaural intensity differences

An experiment was conducted to measure observers' ability to detect time-varying interaural intensity differences (IIDs). In a two-interval forced-choice task, observers discriminated a binaural amplitude modulated (AM) noise in which the modulating sinusoid was interaurally in-phase from the same AM noise in which the modulator was interaurally phase-reversed. The latter stimulus produces a sinusoidally varying IID whose rate and peak IID depend on the frequency (fm) and depth (m) of modulation. The carrier was a narrow-band noise, interaurally uncorrelated, centered at 500, 1000, or 4000 Hz. Presentation level was 75 dB SPL; duration was 1.0 s. For a given fm, m was varied in an adaptive procedure to estimate the depth required for 71% discriminability (mthr). Three of the four observers displayed "low-pass" modulation functions: at 500 Hz, as fm increased from 0-50 Hz, mthr increased from 0.08 (IID = 1.3 dB) to 0.50 (peak IID = 9.5 dB). At 1000 and 4000 Hz observers were more sensitive to IID and the functions (mthr vs fm) were flatter than at 500 Hz. Comparison of these data to previously published data indicates that the binaural system can follow fluctuations in IID more efficiently than it can follow fluctuations in interaural time difference, although there are large individual differences in subjects' capacity to process these two types of binaural cues.