First-order Intervals Research Articles

Behavioral and physiological correlates of temporal pitch perception in electric and acoustic hearing

In the "4-6" condition of experiment 1, normal-hearing (NH) listeners compared the pitch of a bandpass-filtered pulse train, whose inter-pulse intervals (IPIs) alternated between 4 and 6 ms, to that of isochronous pulse trains. Consistent with previous results obtained at a lower signal level, the pitch of the 4-6 stimulus corresponded to that of an isochronous pulse train having a period of 5.7 ms-longer than the mean IPI of 5 ms. In other conditions the IPI alternated between 3.5-5.5 and 4.5-6.5 ms. Experiment 2 was similar but presented electric pulse trains to one channel of a cochlear implant. In both cases, as overall IPI increased, the pitch of the alternating-interval stimulus approached that of an isochronous train having a period equal to the mean IPI. Experiment 3 measured compound action potentials (CAPs) to alternating-interval stimuli in guinea pigs and in NH listeners. The CAPs to pulses occurring after 4-ms intervals were smaller than responses to pulses occurring after 6-ms intervals, resulting in a modulated pattern that was independent of overall level. The results are compared to the predictions of a simple model incorporating auditory-nerve (AN) refractoriness, and where pitch is estimated from first-order intervals in the AN response.

Pitch strength of regular-interval click trains with different length “runs” of regular intervals

Click trains were generated with first- and second-order statistics following Kaernbach and Demany [J. Acoust. Soc. Am. 104, 2298-2306 (1998)]. First-order intervals are between successive clicks, while second-order intervals are those between every other click. Click trains were generated with a repeating alternation of fixed and random intervals which produce a pitch at the reciprocal of the duration of the fixed interval. The intervals were then randomly shuffled and compared to the unshuffled, alternating click trains in pitch-strength comparison experiments. In almost all comparisons for the first-order interval stimuli, the shuffled-interval click trains had a stronger pitch strength than the unshuffled-interval click trains. The shuffled-interval click trains only produced stronger pitches for second-order interval stimuli when the click trains were unfiltered. Several experimental conditions and an analysis of runs of regular and random intervals in these click trains suggest that the auditory system is sensitive to runs of regular intervals in a stimulus that contains a mix of regular and random intervals. These results indicate that fine-structure regularity plays a more important role in pitch perception than randomness, and that the long-term autocorrelation function or spectra of these click trains are not good predictors of pitch strength.

Discrimination of first- and second-order regular intervals from random intervals as a function of high-pass filter cutoff frequency

This study extends the work of Kaernbach and Demany [J. Acoust. Soc. Am. 104, 2998–2306 (1998)] in which regular interval stimuli (RIS) click trains with first-order intervals could be discriminated from random-interval click trains, but RIS with second-order intervals could not. Kaernbach and Demany concluded that their results cast doubt on autocorrelation as a method of analysis for such stimuli. The present study investigated the same stimuli, but for a variety of filter conditions. The results suggest that while RIS click trains with first-order intervals are more easily discriminated from random-interval stimuli than second-order interval RIS click trains, discrimination based on second-order intervals is possible except when the stimuli are high-pass filtered above 8 kHz, i.e., above the spectral region of phase locking.

Limits of rhythm perception.

To what extent are listeners sensitive to the time intervals separating non-consecutive events in sound sequences? The subjects of Experiment 1 were presented with sequences of 20 identical tones in which the 10 odd-numbered tones or the 10 even-numbered tones made up an isochronous sub-sequence (with a periodicity of 0.5-1 s) whereas the other tones, acting as distractors, occurred at random moments. Such sequences appeared to be very difficult to discriminate from sequences without any timing regularity, which revealed a lack of perceptual sensitivity to their "second-order" intervals. Experiment 2 employed repetitive sequences in which the first-order intervals (separating consecutive tones) took two possible values, forming a ratio that subjects had to classify as larger or smaller than 2. The results of this experiment suggest that subjects were able to make use of second-order intervals in their task, but mainly due to the predictable nature of the sequences; the relative positions of subjective accents (Povel & Essens, 1985) had no significant effect on performance. It is concluded that the perception of subtle timing details in "ordinary" music may rest on nothing more than a sensitivity to the relations between first-order intervals (within a given auditory stream).

Temporal representation of iterated rippled noise as a function of delay and sound level in the ventral cochlear nucleus.

The discharge patterns of single units in the ventral cochlear nucleus (VCN) of anesthetized guinea pigs were examined in response to iterated rippled noise (IRN) as a function of the IRN delay (which determines the IRN pitch) and the IRN sound level. Delays were varied over five octaves in half-octave steps, and sound levels were varied over a 30- or 50-dB range in steps of 5 dB. Neural responses were analyzed in terms of first-order and all-order inter-spike intervals (ISIs). The IRN quasi-periodicity was preserved in the all-order ISIs for most units independent of unit type or best frequency (BF). A deterioration of the temporal all-order code was found, however, when the neural response was influenced by inhibition. The IRN quasi-periodicity was also preserved in first-order ISIs for a limited range of IRN delays and levels. Sustained Chopper units (CS) in the VCN responded with very regular ISIs when the IRN delay corresponded to the unit's chopping period; i.e., the unit showed an increased proportion of intervals corresponding to the IRN delay (interval enhancement) relative to an equal-level, white-noise stimulation. This interval enhancement has a band-pass characteristic with a peak corresponding to the chopping period. Moreover, for CS units in rate saturation, the chopping period, and thus the interval enhancement to the IRN, did not vary with level. Units classified as onset-chopper also show a band-pass interval enhancement to the IRN stimuli; however, they show more level-dependent changes than CS units. Primary-like (PL) units also show level-dependent changes in their ability to code the IRN pitch in first-order intervals. The range of delays where PL units showed interval enhancement was broader and extended to shorter delays. Based on these findings, it is suggested that CS units may play an important role in pitch processing in that they transform a higher-order interval code into a first-order interval place code. Their limited dynamic range together with the preservation of the temporal stimulus features in saturation may serve as a physiological basis for the perceived level independence of pitch.

Lateral septal neuronal firing rate increases during proestrus-estrus in the rat

Neuronal activity of the lateral septal nucleus (LSN) is related to motivational and hedonic behavior. Even though some changes in mood and anxiety during proestrus and pregnancy have been reported, the possible changes in the neuronal activity of the LSN through the phases of the estrous cycle are unknown. In the present study we explored the neuronal activity from the LSN using glass micropipettes (NaCl 1 M, and Evans blue 2.5%; 3–8 MΩ) in 30 urethane (1 g/kg) anesthetized Wistar rats. Analysis of data included a total of 88 single-unit extracellular recordings taken from the LSN during proestrus ( n = 22), estrus ( n = 23), diestrus ( n = 22), and metestrus ( n = 21). The highest values of firing rate were found in proestrus, and the lowest in metestrus, F(3, 84) = 3.78, p < 0.01. During estrous cycles, in the phase characterized by high plasma levels of estradiol and progesterone, i.e., proestrus–estrus, the neurons from the dorsal aspect of the LSN fired at significantly ( p < 0.05) higher frequencies, shorter first-order intervals and a lower coefficient of variation than those in the phase characterized by lower levels of estradiol and progesterone (metestrus–diestrus). In another group of rats ( n = 12), immobility in the forced-swim test was assessed. Consistently, a longer latency ( p < 0.05) for the first period of immobility and a nonsignificant trend to a lowered total time in immobility were found in proestrus and estrus. It is concluded that the higher firing rate in neurons from the dorsal aspect of the LSN during proestrus–estrus, may be associated with an increased motivation to escape from a stressful situation.

Effect of stimulation on burst firing in cat primary auditory cortex

1. Neural activity was recorded extracellularly with two independent microelectrodes aligned in parallel and advanced perpendicular to isofrequency sheets in cat primary auditory cortex. Multiunit activity was separated into single-unit spike trains using a maximum variance spike sorting algorithm. Only units that demonstrated a high quality of sorting and a minimum spontaneous firing rate of 0.2 spikes/s were considered for analysis. The primary aim of this study was to describe the effect of periodic click train and broadband noise stimulation on short-time-scale (< or = 50 ms) bursts in the spike trains of single auditory cortical units and to determine whether stimulation influenced the occurrence, spike count, and/or temporal structure of burst firing relative to a spontaneous baseline. 2. Extracellular recordings were made in 20 juvenile and adult cats from 69 single auditory cortical units during click train stimulation and silence, and from 30 single units during noise stimulation and in silence. In an additional 15 single units the effect of both click train and noise stimulation was investigated. The incidence, spike count, and temporal structure of short-time-scale burst firing in the first 100 ms following stimulus presentation was compared with burst firing in the period starting 500 ms after stimulus presentation and with spontaneous burst firing. In addition, the serial dependence of interspike intervals within a burst was tested during periods of stimulation. 3. Burst firing was present in the stimulation, poststimulation, and spontaneous conditions. Longer bursts (consisting of > or = 3 spikes) were more commonly observed in the poststimulation and spontaneous conditions than in the stimulation condition. This effect was most pronounced during click stimulation. A period of elevated firing activity was present in a subset of units 0.5-1.5 s after stimulus presentation, indicating prolonged effects of stimulation on single-unit firing behavior. 4. For both stimuli, the proportion of single-unit responses composed of bursts was significantly greater in poststimulation and spontaneous periods than during stimulation. Burst rate was higher in post-click-train stimulation and spontaneous periods than during periods of click stimulation. The isolated spike rate was significantly higher during periods of noise and click stimulation than in the poststimulation and spontaneous periods. 5. An examination of the autocorrelograms and higher-order interspike interval histograms of single-unit responses during click train stimulation indicated that 25% of single-unit spike trains contained an excess of brief first-order intervals and 14% of spike trains contained a shortage of long higher-order interspike intervals relative to a spontaneous baseline. During noise stimulation, 10% of single-unit responses contained an excess of short intervals relative to baseline. Interspike intervals of short-duration bursts were not serially dependent during periods of stimulation. 6. A comparison of the autocorrelograms and higher-order interval histograms of single-unit responses in the poststimulation and spontaneous conditions indicated that 20% of single-unit spike trains contained an excess of short first-, second-, and third-order intervals following stimulation. This subgroups of single units could not be distinguished on the basis of the age of the animal or the depth at which the recording was made. 7. The low incidence of burst firing during stimulation opposes the view that bursts serve as a mechanism to emphasize or amplify particular stimulus-related responses in the presence of ongoing spontaneous activity in the primary auditory cortex. Moreover, there is little evidence to support the notion that brief bursts represent neural codes, because intraburst intervals are not serially dependent. It is suggested that pyramidal burst firing may be an effective way to evoke postsynaptic firing in inhibitory interneurons and subsequ