Tonal frequency shifts and gaps in acoustic stimulation as reflex-modifying events

When a relatively weak signal, such as a mild tone, precedes an intense reflex-eliciting stimulus by an appropriate interval (about 100 msec), the amplitude of the elicited reaction is often reduced. It was found that in student volunteers a brief gap in a steady pure tone that occurred 150 msec prior to a mild tap to the glabella (the flat region between the eyebrows) could inhibit the eyeblink elicited by the tap. It was also found that a shift in tonal frequency across a gap in a tone was more inhibitory than a gap with no frequency shift, but it was no more inhibitory than the onset of the short second tone alone. The final study determined the minimum amount of frequency shift required to produce an additional inhibitory effect above that of a gap alone. The findings are discussed in terms of various aspects of sensory processing.     

Startle reflex inhibition in the rat: its persistence after extended repetition of the inhibitory stimulus

Startle reflexes to intense sound bursts are inhibited by weak stimuli that briefly precede their elicitation. In three experiments the startle stimulus (a 110-dB SPL tone burst) was presented 100 ms after the final link in a train of stimuli, the length of the train varying from 1 to 1,000, its repetition rate varying from 1 per s to 10 per s, and its constituents being 40 dB or 50 dB white noise bursts of 25 ms duration. Inhibition was invariant across train length and repetition rate. In a final experiment the startle stimulus was presented a variable interval after the final link, from 40 ms to 1280 ms, with 1 or 100 noise bursts (50 dB) in the train. Inhibition developed more rapidly following the last member of the 100-stimulus train, suggestive of a "priming" or sensitization effect of stimulus repetition, but its overall strength and subsequent rate of decay were not different in the two conditions. The general persistence of inhibition following these extended series of stimuli reveals that reflex inhibition must be the outcome of a fixed and obligatory process associated with sensory input.     

Temporal specificity of fear conditioning: effects of different conditioned stimulus-unconditioned stimulus intervals on the fear-potentiated startle effect

Separate groups of rats were given 30 pairings of a light (conditioned stimulus, CS) and 500-ms shock (unconditioned stimulus, US) at CS-US intervals of 0, 25, 50, 100, 200, 800, 3,200, 12,800, or 51,200 ms. Other groups had lights and shocks inconsistently paired. The startle reflex was elicited 2-4 days later with a noise burst alone or 25-51,200 ms after light onset. After CS-US pairings over a wide range of intervals (25-51,200 ms), startle was potentiated in testing, sometimes as rapidly as 50 ms after light onset. Magnitude of potentiation and resistance to extinction were generally greater with longer CS-US intervals. In several groups, potentiation was maximal at a test interval that matched the CS-US interval used in training. This temporal specificity sharpened with increasing numbers of training trials but even occurred with a single training trial in which a 51,200-ms CS-US interval was used. The data indicate that even during simple fear conditioning, animals rapidly learn a temporal CS-US relationship. This has important implications for understanding the neural mechanisms of fear conditioning.     

Echoic memory in the rat: effects of inspection time, retention interval, and the spectral composition of masking noise.

Memory for tones (1100 vs. 2330 Hz) was studied in 4 rats (Rattus norvegicus), as affected by the durations of both target tones (30 to 620 ms) and noise-filled retention intervals (0 to 480 ms). With a 0-ms delay, performance was near asymptotic with the 30-ms tone, but the memory of this brief tone suffered a massive decrement at retention intervals as brief as 60 ms; in contrast, memory for the 340-ms tone was stable for at least 240 ms. If the retention interval was filled by band-stop noise (with targets presented in the spectral gap), then the rat's memory for brief tones was superior to that obtained with the standard broad-band noise filler, and band-stop noise was better than a band-pass noise that had the tones embedded in the region of its spectral energy. These findings are consistent with the hypotheses that auditory memory in the rat consists of a transient sensorylike echoic store and a short-term store more resistant to the effects of retroactive interference.     

Spectral frequency and the modulation of the acoustic startle reflex by background noise

The rat's (Long-Evans) acoustic startle reflex to a high-frequency tone burst (10.5 kHz) was depressed by intense high-frequency band-pass noise (8-16 kHz) but enhanced by low frequency noise (1-2 kHz). However, contrary to the hypothesis that the depression of startle in intense background noise is produced by sensory masking, the reflex to a low-frequency tone burst (at 1 kHz) was depressed by both high- and low-frequency band-pass noise. Two additional hypotheses are offered to supplement sensory masking in order to explain the asymmetry in these data. The first is that the intratympanic reflex, which acts as a high pass filter on acoustic input, is elicited in intense backgrounds. The second is that acoustic startle reflexes elicited by intense low-frequency tones are in part elicited by their high-frequency distortion products and that these distortion products are then masked by high-frequency background noise.     

The startle response and auditory temporal summation in neonates

The present study assessed temporal summation of transient and sustained stimuli in the startle eyeblink response system in neonates during quiet sleep. Subjects received 100-dB(A), fast-rising broadband noise bursts of two types: (a) single stimuli varying in duration from 20 to 100 ms and (b) pairs of 3-ms bursts presented at interpulse intervals corresponding to the single-stimulus durations. In addition, a single 3-ms pulse was used as an anchor point for both stimulus types. For startle amplitude, single stimuli were more effective than were paired stimuli, but the temporal summation functions were similar for the two types of stimuli. Response amplitude increased as stimulus duration/interval increased to 50 ms, but not beyond. For startle probability, temporal summation was similar for single and paired stimuli at 20 ms. Pairs of pulses were equally effective at 20, 35, and 50 ms, beyond which the second pulse was not effective. Increasing the duration of single stimuli from 20 to 35 ms resulted in increased probability, illustrating a contribution of sustained summation beyond that of transient summation. Response latency was generally greater for paired than for single stimuli. The results suggest that temporal summation of brief stimuli is deficient in the neonate. These data were compared to adult data from an analogous study, and suggest that the transient system is immature in infants, and that this immaturity is expressed differently by startle amplitude, probability, and latency.     

Changes in the magnitude of the eyeblink startle response during habituation of sexual arousal

Modulation of the startle response was used to examine emotional processing of sexual stimulation across trials within a session. Eyeblink startle was elicited by a probe (burst of intense white noise) presented intermittently while men were viewing an erotic film segment. Repeated display of the film segment resulted in a progressive decrease in sexual arousal. Habituation of sexual arousal was accompanied by a reduction over trials in the extent the men felt absorbed when viewing the erotic stimulus and by an increase over trials in the magnitude of the eyeblink startle response. Replacing the familiar stimulus by a novel erotic stimulus increased in sexual arousal and absorption and reduced startle (novelty effect), while dishabituation was evident for all three response measures when the familiar stimulus was reintroduced. This pattern of results indicates that with repeated presentation an erotic stimulus is experienced not only as less sexually arousing but also as less appetitive and absorbing. The question of whether habituation of sexual arousal is mediated by changes in attentional and affective processing over trials is discussed, as are clinical contexts in which eyeblink startle can be used in studying aspects of sexual functioning.     

Auditory change detection: simple sounds are not memorized better than complex sounds

Previous research has shown that the detectability of a local change in a visual image is essentially independent of the complexity of the image when the interstimulus interval (ISI) is very short, but is limited by a low-capacity memory system when the ISI exceeds 100 ms. In the study reported here, listeners made same/different judgments on pairs of successive "chords" (sums of pure tones with random frequencies). The change to be detected was always a frequency shift in one of the tones, and which tone would change was unpredictable. Performance worsened as the number of tones increased, but this effect was not larger for 2-s ISIs than for 0-ms ISIs. Similar results were obtained when a chord was followed by a single tone that had to be judged as higher or lower than the closest component of the chord. Overall, our data suggest that change detection is based on different mechanisms in audition and vision.     

Specificity and savings of behavioural habituation over a series of intra- and inter-modal stimuli

A series of four tone stimuli, differing either in tonal frequency or in melody, was presented at the rate of one presentation per day to a group of rats. Habituation was specific to the frequency of simple auditory stimuli and to the melody of patterned stimuli, as shown both by the distraction to stimulus change, and by the trials to habituate to the new stimulus. Savings in habituation occurred when a series of tone stimuli was presented, but was not found for a series of inter-modal stimuli.     

Potentiation of acoustic startle behavior in the rat (Rattus norvegicus) at the onset of darkness

In Experiment 1 (N = 8), a 20-ms light pulse, given at various times before a noise burst, inhibited reflex expression with a single trough at a lead time of 70 ms, whereas a dark pulse facilitated the reflex with two peaks at 40 and 160 ms. In Experiment 2 (N = 18) facilitation by dark onset had a single peak, and inhibition by light onset a single trough; thus, the double peak of the dark pulse may result because inhibition from light onset at the end of the dark pulse was briefly impressed on the facilitatory effect of dark onset. In Experiment 3 (N = 12), diazepam (2.5 mg/kg, but not 1 mg/kg) eliminated dark facilitation but not light inhibition. These diazepam data reveal a basic similarity, perhaps identity, of the mechanisms responsible for the effect of dark onset and those producing reflex facilitation by Pavlovian fear conditioning and prolonged background noise, because all are moderated by a GABAergic system.     

The role of small changes in the acoustic environment in modifying the startle reflex.

When either the intensity or frequency spectrum of an approximately 70-db. SPL narrow-band noise was abruptly changed by a small amount, the rat's response to a startle stimulus presented 64 msec later was inhibited. When similar small frequency changes preceded the startle stimulus by ony 5 msec, the latency of the startle response was reduced, but even relatively large changes in intensity of the antecedent stimulus had no effect on response latency. These findings provide added support for the generalization that the neural processes associated with startle are engaged by small changes in the auditory environment. They also point to a measure of separation between the processes responsible for inhibition and those responsible for latency shift.     

Avoidance learning facilitates temporal processing in the primary auditory cortex

The primary auditory cortex is now known to be involved in learning and memory, as well as auditory perception. For example, spectral tuning often shifts toward or to the frequency of the conditioned stimulus during associative learning. As previous research has focused on tonal frequency, less is known about how learning might alter temporal parameters of response in the auditory cortex. This study addressed the effects of learning on the fidelity of temporal processing. Adult male rats were trained to avoid shock that was signaled by an 8.0 kHz tone. A novel control group received non-contingent tone and shock with shock probability decreasing over days to match the reduced number of shocks received by the avoidance group as they mastered the task. An untrained (nai ve) group served as a baseline. Following training, neuronal responses to white noise and a broad spectrum of tones were determined across the primary auditory cortex in a terminal experiment with subjects under general anesthesia. Avoidance conditioning significantly improved the precision of spike-timing: the coefficient of variation of 1st spike latency was significantly reduced in avoidance animals compared to controls and nai ves, both for tones and for noise. Additionally, avoidance learning was accompanied by a reduction of the latency of peak response, by 2.0-2.5 ms relative to nai ves and approximately 1.0 ms relative to controls. The shock-matched controls also exhibited significantly shorter peak latency of response than nai ves, demonstrating the importance of this non-avoidance control. Plasticity of temporal processing showed no evidence of frequency specificity and developed independently of the non-temporal parameters magnitude of response, frequency tuning and neural threshold, none of which were facilitated. The facilitation of temporal processing suggests that avoidance learning may increase synaptic strength either within the auditory cortex, in the subcortical auditory system, or both.     

Modification of the rat's acoustic startle response by antecedent visual stimulation.

Male hooded and albino rats were exposed to a light flash followed at various temporal intervals by a startle-eliciting 117 db. (re 20 muN/m2) burst of white noise. The visual stimulus engendered startle response inhibition (maximally when the lead time was 64-250 msec) as well as startle response latency reduction (maximally when the lead time was 2-8 msec). The temporal functions for the effects of visual stimuli paralleled those previously reported for startle modification by acoustic events. Further study revealed that, given optimal lead times, inhibition is produced reliably by weaker visual stimuli (3 X 10-6 cd-sec/cm2) than latency reduction (3 X 10-4 cd-sec/cm2). This differential sensitivity to visual stimuli is also analogous to previously reported findings for events in the acoustic environment. It reveals that the neural mechanisms that mediate latency reduction and inhibition can be engaged by either acoustic or visual stimulation.     

Auditory masking-level differences in the cat.

Four cats were trained to avoid shock by responding to the intermittent occurrence of 1-kHz tone pulses at one ear, while a continual train of noise pulses was simultaneously presented either to the signal ear alone or to both ears. Using the masked threshold levels determined with monaural noise as a reference, the amount of unmasking produced by the addition of noise to the nonsignal ear was measured. Significantly lower tonal detection thresholds were observed when noise equal in intensity to that at the signal ear was added to the nonsignal ear. Additional unmasking occurred when the intensity of the noise at the latter ear was raised to a level 10 db. higher than that at the signal ear.     

Similar effects of attention directed to acoustic and tactile stimuli on prepulse inhibition of acoustic startle

Prepulse inhibition (PPI) is assumed to index automatic and controlled processing. In three experiments (n= 32, 22, and 30) participants were asked to judge the duration of a prepulse in comparison with a stimulus presented 4000 ms before the prepulse. A distracter was presented simultaneously with the prepulse to increase the cognitive demands of the task. PPI was assessed at stimulus onset asynchronies (SOAs) of 30-150 ms, and 420 ms. The prepulse was either a tone (60 dB) or a tactile stimulus (21 kPa), and startle was elicited by 95 dB white noise. Directing attention to the prepulse increased PPI at SOAs of 60 ms and longer in all experiments, but the sensory modality to which attention was directed played only a minor role. We conclude that directing attention to both acoustic and tactile prepulses increased PPI.     

Age-related changes in auditory temporal processing

Two tone bursts separated by a silent interval and imbedded in a background white noise were presented to elderly subjects (M age = 71.3 years) and young subjects (M age = 22.2 years). Subjects were required to judge when the two tone bursts fused perceptually by adjusting the duration of tone-one. The bursts were separated by six discrete interstimulus intervals of 4, 8, 16, 24, 32, or 40 ms. The tone-two burst was held constant at 100 ms. Fusion point was defined as that critical tone-one duration at which the two tones fused (were perceived as one). Elderly subjects reached fusion point at longer critical tone-one durations than young subjects at each interval tested. The function relating tone-one duration and interval conformed to an exponential curve and is discussed with respect to an exponential decay model of the inhibitory interactions of neural systems responding to onsets and offsets of sensory events.     

The extent of active processing of a long-duration stimulus modulates the scalp-recorded sustained potential

A long-duration stimulus will elicit a negative sustained potential (SP) that is maximum in amplitude over fronto-central areas of the scalp. This study examines how the duration of active attentional processing of the stimulus might also elicit a nonsensory contingent negative variation (CNV) that overlaps and summates to the SP. Subjects were presented with either a low- or high-pitched 1.4s duration stimulus. In different conditions, a 20 ms gap occurred either 300 or 1300 ms after stimulus onset in half the stimuli. The subject's task was to button press upon detection of the gap in the low-pitched stimulus. The subject was not required to respond to the gap occurring in the high-pitched tone. In a separate Ignore condition, subjects ignored all stimuli and read a book. In the Ignore condition, a SP was apparent to the long-duration stimuli. In the Attend condition, a large amplitude slow wave, probably the CNV, overlapped and summated with the SP. The extent of the overlapping CNV was dependent on whether the gap occurred early or late. The CNV was not apparent following presentation of the high-pitched tone, when the gap did not need to be detected. Consistent with previous studies, a SP was thus elicited even when active processing of the stimulus was not required. The morphology of the scalp-recorded SP was, nevertheless, much modified by an overlapping and summating CNV, depending on the extent to which active processing of the long-duration stimulus was required.     

Repetition priming effects from attended vs. ignored single words in a semantic categorization task

The present research examines priming effects from a centrally presented single-prime word to which participants were instructed to either attend or ignore. The prime word was followed by a single central target word to which participants made a semantic categorization (animate vs. inanimate) task. The main variables manipulated across experiments were attentional instructions (attend vs. ignore the prime word), presentation duration of the prime word (20, 50, 80 or 100 ms), prime-target stimulus onset asynchrony (SOA; 300 vs. 800 ms), and temporal presentation of instructions (before vs. after the prime word). The results showed (a) a consistent interaction between attentional instructions and repetition priming and (b) a qualitatively different ignored priming pattern as a function of prime duration: reduced positive priming (relative to the attend instruction) for prime exposures of 80 and 100 ms, and reliable negative priming for the shorter prime exposures of 20 and 50 ms. In addition (c), the differential priming pattern for attend and ignore trials was observed at a prime-target SOA of 800 ms (but not at a shorter 300-ms SOA) and only when instructions were presented before the prime word. Methodological and theoretical implications of the present findings for the extant negative priming literature are discussed.     

Responsiveness of Western adults to pitch-distributional information in melodic sequences

Responsiveness of musically trained and untrained adults to pitch-distributional information in melodic contexts was assessed. In Experiment 1, melodic contexts were pure-tone sequences, generated from either a diatonic or one of four nondiatonic tonesets, in which pitch-distributional information was manipulated by variation of the relative frequency of occurrence of tones from the toneset. Both the assignment of relative frequency of occurrence to tones and the construction of the (fixed) temporal order of tones within the sequences contravened the conventions of western tonal music. A probe-tone technique was employed. Each presentation of a sequence was followed by a probe tone, one of the 12 chromatic notes within the octave. Listeners rated the goodness of musical fit of the probe tone to the sequence. Probe-tone ratings were significantly related to frequency of occurrence of the probe tone in the sequence for both trained and untrained listeners. In addition, probe-tone ratings decreased as the pitch distance between the probe tone and the final tone of the sequence increased. For musically trained listeners, probe-tone ratings for diatonic sequences tended also to reflect the influence of an internalized tonal schema. Experiment 2 demonstrated that the temporal location of tones in the sequences could not alone account for the effect of frequency of occurrence in Experiment 1. Experiment 3 tested musically untrained listeners under the conditions of Experiment 1, with the exception that the temporal order of tones in each sequence was randomized across trials. The effect of frequency of occurrence found in Experiment 1 was replicated and strengthened.     

The sensory nature of knowledge: Sensory priming effects in semantic categorization

The aim of the present study was to show the perceptual nature of conceptual knowledge by using a priming paradigm that excluded an interpretation exclusively in terms of amodal representation. This paradigm was divided into two phases. The first phase consisted in learning a systematic association between a geometrical shape and a white noise. The second phase consisted of a short-term priming paradigm in which a primed shape (either associated or not with a sound in the first phase) preceded a picture of an object, which the participants had to categorize as representing a large or a small object. The objects were chosen in such a way that their principal function either was associated with the production of noise (“noisy” target) or was not typically associated the production of noise (“silent” target). The stimulus onset asynchrony (SOA) between the prime and the target was 100 ms or 500 ms. The results revealed an interference effect with a 100-ms SOA and a facilitatory effect with a 500-ms SOA for the noisy targets only. We interpreted the interference effect obtained at the 100-ms SOA as the result of an overlap between the components reactivated by the sound prime and those activated by the processing of the noisy target. At an SOA of 500 ms, there was no temporal overlap. The observed facilitatory effect was explained by the preactivation of auditory areas by the sound prime, thus facilitating the categorization of the noisy targets only.