Temporal integration of acoustic and cutaneous stimuli shown in the blink reflex

Temporal integration of pairs of brief blink-eliciting acoustic and cutaneous stimuli was investigated to determine if there was integration of stimuli from different modalities. Reflexes elicited by a tone burst or by a brief electrical shock to the supraorbital nerve followed by a second tone burst or shock at short stimulus onset asynchronies (SOAs) were larger and faster than control reflexes elicited by a single stimulus identical to the lead stimulus of the stimulus pairs. Reflex amplitude was augmented at longer SOAs where there was no effect on latency. Temporal integration was evident for all stimulus pairs, showing that it is due, at least in part, to processes that occur outside specific sensory pathways. Heterogeneous stimulus pairs produced greater reflex enhancement than did homogeneous stimulus pairs. This finding was examined further in Experiment 2, which showed that reflex enhancement with pairs of acoustic pulses was unaffected by the frequency of the second stimulus, suggesting that sensory masking was not acting to suppress reflex expression with acoustic pulse pairs. Integration of reflexogenic acoustic stimuli shown in the blink reflex is restricted to shorter intervals than is integration of acoustic stimuli shown by psychophysical procedures, suggesting that the two methods reflect different aspects of stimulus processing. Integration of reflexogenic stimuli may result from summation of activity associated more directly with reflex expression than with perceptual awareness.     

Modulation of the acoustic startle reflex in humans in the absence of anticipatory changes in the middle ear reflex

If a weak tone precedes an intense tone, then the acoustic startle eyeblink reflex elicited by the stronger stimulus is inhibited. It has been suggested that the leading stimulus gives rise to a protective middle ear reflex that attenuates the effective intensity of the second. This hypothesis was tested and disproved. In seven subjects intense tone bursts sufficient to elicit both intratympanic and eyeblink responses were presented sometimes alone and sometimes preceded at various lead times (25 to 400 msec) by a weak tone. The weak tone inhibited the amplitude of the eye blink to the strong tone, maximally at intervals of 100 to 200 msec, but was never seen to produce any of the anticipatory impedance changes that would be characteristic of middle ear reflex activity during the interval between the two stimuli.     

Acoustic startle threshold of the albino rat (Rattus norvegicus)

The startle threshold of the albino Sprague-Dawley rat runs parallel to the curve of the hearing threshold. The difference between the startle and hearing threshold is 87 dB (SPL) at a background noise level of 75 dB (SPL). At 110 dB (SPL), the threshold has a range from 2 kHz to 50 kHz with a minimum at 10 kHz and a second minimum at 40 kHz. Amplitude and latency of the startle response are not only dependent on the sensation level of the acoustic stimulus but also on the frequency. At threshold, only the head movement component of the startle response is elicited.     

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.     

Reflex modification in the rat: the inhibitory effects of intensity and frequency changes in steady tones

In laboratory rats (as in humans) a low-intensity tone that precedes a high-intensity burst of noise by approximately 100 ms can reduce the amplitude of the startle reaction elicited by the burst of noise. A series of four experiments with rats investigated the relation between the inhibitory effects of tonal frequency change and the length of the silent period (gap) preceding it. The major findings were the following: (a) A gap in an otherwise continuous pure tone inhibited startle when the gap occurred approximately 100 ms prior to the noise burst. (b) Although an increase in gap duration increased the inhibition afforded by the gap, the maximum inhibition was yielded by gaps of 100 ms and greater; this maximum was equivalent to the inhibition yielded by the presentation of a postgap tone alone. (c) A shift in tonal frequency across a 10-ms gap yielded more inhibition than did the same gap with no frequency shift; again the shift yielded equivalent inhibition to the presentation of the postgap tone alone. (d) An increase in the frequency shift increased inhibition when the shift occurred across a 10-ms gap, but not when the shift occurred across a 100-ms gap.     

Spatial-frequency masking with briefly pulsed patterns

Spatial-frequency masking was studied with briefly pulsed (25 ms) vertical gratings. The mask was a noise grating, and the test pattern was a sinusoidal grating. A low-frequency band of noise masked a low- but not high-spatial-frequency test grating when the patterns were presented simultaneously. A high-frequency band of noise did not mask a low-frequency test grating when the patterns were presented simultaneously or when the mask was presented after the test pattern (backward masking). Masking was, however, observed when the mask or test pattern was of sufficiently high contrast so that the stimuli had nonlinear distortion and thus produced DC shifts of the field luminance.     

Directed attention influences the modification of startle reflex probability

The present study was designed to investigate the effect of directed attention on elicitation and modification of the startle reflex. 30 adult human subjects received 90 dB(A) broadband noise startle stimuli either alone or preceded by a 60 dB(A) prepulse (either 2000-Hz tone, 1000-Hz tone, or broadband noise). Subjects were instructed to attend to one of the three prepulses during half the trials and to ignore all stimuli during the rest of the trials. The probability of responding while attending to a prepulse was significantly lower than the probability of responding while ignoring the prepulses. Responding to the prepulses was also more probable while subjects were attending to the prepulses, and these effects were more pronounced for tone than for noise prepulses. These results suggest that directed attention can influence the probability of the startle reflex without influencing startle amplitude or latency.     

Modality-repetition and attentional effects on reflex blinking in infants and adults

The ability of a brief prepulse to inhibit reflexively elicited startle is a robust phenomenon in adults of several species but is weak or absent in infants. The possibility that afferent processing of the prepulse is inadequate in infants was tested by assessing the integrity of two other types of reflex modulation that occur in modality-specific paths, that is, attenuation by repetition of same-modality stimuli and enhancement by modality-selective attention. The reflex, measured by electromyographic activity of the muscle controlling blink, was elicited by intense light flashes or noise bursts preceded by brief acoustic or visual prepulses and delivered during attention-directing acoustic or visual foregrounds that evoked cardiac deceleration. Modality-repetition effects were evidenced by smaller peak blink magnitude and longer latency of blinking to same than to different modality pairs and did not differ as a function of age (4-month-old infants or college students). Attention effects were also seen in both infants and adults and, in accord with previous findings, were evident in magnitude for infants and latency for adults. Thus, immaturity in paths mediating these effects could not explain delayed development of prepulse inhibition. Other possibilities include delayed maturation of extrinsic inhibition or of transient-processing systems.     

Sources of auditory masking in infants: distraction effects.

Previous work has demonstrated that infants' thresholds for a pure tone are elevated by a masker more than would be predicted from their critical bandwidths. The present studies explored the nature of this additional masking. In Experiment 1, detection thresholds of 6-month-old infants and of adults for a 1-kHz tone were estimated under three conditions: in quiet, in the presence of a 4- to 10-kHz bandpass noise at 40 dB SPL, and in the presence of the same noise at 50 dB SPL. The noise was gated on at the beginning of each trial. Adult thresholds were the same in all three conditions, indicating that little or no sensory masking took place in the presence of the noise. Infant thresholds were about 10 dB higher in the presence of the noise. We term this effect distraction masking. In Experiment 2, the effect of gating the noise on at trial onset was examined. Thresholds for the same tone were estimated in quiet and in the presence of the band-pass noise at 40 dB SPL, but the noise was presented continuously during the session. Under these conditions, distraction masking was still observed for infants. These findings suggest that a masker can have nonsensory effects on infants' performance in a psychoacoustic task.     

A signal detection theory analysis of gap detection in the rat

An innocuous sensory event (a prestimulus) briefly preceding a startle-eliciting stimulus (SES) reduces the amplitude of the elicited reflex. This study used signal detection theory (SDT) techniques to quantify the effects of gaps (pauses in otherwise continuous noise) on the rat’s acoustic startle reflex. Sixteen rats were given four identical test sessions consisting of the randomized presentation of 150 trials of the SES alone and 150 trials of a gap-and-SES combination. Gap duration (1, 2, 4, and 8 msec) varied between sessions. Data analyses based on amplitude, difference scores, percentage scores, and SDT techniques identified similar patterns. The three longest gaps, but not the shortest, were reliably detected, and differences among these three were identified with percentage and SDT analyses. Analyses of amplitude changes over test sessions yielded different patterns for each measure. The results demonstrate that an SDT analysis is a sensitive index of prestimulus effects.     

Parameters affecting gap detection in the rat

The present research used a startle amplitude reduction paradigm to investigate the ability of the rat’s auditory system to track rapidly changing acoustic transients. Specifically examined was the ability of brief gaps in otherwise continuous noise to reduce the amplitude of a subsequently elicited acoustic startle reflex. The duration of the gap, time between gap offsetand startle elicitation (the interstimulus interval or ISI), and rise-fall characteristics of the gap were systematically varied. Consistent with previous research, gaps reliably reduced startle amplitude. Gaps 2 msec long were reliably detected, and a 50-msec ISI resulted in the greatest amplitude reduction. Gaps presented at short ISIs produced amplitude reduction that followed a different time course than did gaps presented at longer ISIs. These results may reflect differences in the length oftime available for the processing of the stimulus and may involve two different processes.     

Effects of caffeine on the trigeminal blink reflex

The acoustic startle and trigeminal blink reflexes share the same motor output. Since caffeine has been shown to augment the startle reflex, it was proposed that caffeine would also increase the trigeminal blink reflex. In 6 humans, the effects of caffeine (100 mg) on the trigeminal blink reflex were investigated. Reflex blinks were elicited by stimulation of the supraorbital branch of the trigeminal nerve. Following ingestion of caffeinated coffee, reflex blinks increased in amplitude and duration and occurred at a shorter latency than reflex blinks following ingestion of decaffeinated coffee. Since the blink reflex is a brainstem reflex, these results suggest that the psychomotor effects of caffeine facilitate brainstem processing.     

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.     

Asymmetry of masking between noise and tone

A pure tone was used to mask narrow and wide bands of noise centered on the frequency of the tone. In a given experimental session, the sound-pressure level (SPL) of the tone was held constant and loudness balances were obtained between a masked and unmasked noise band of equal width. These results are compared to earlier measures of the partial masking of tone by noise. The comparison shows that noise masks a tone more effectively than the tone masks the noise. Although the effect of the tone on a critical band of noise is greater than its effect on either an octave-band noise or wide-band noise, it is considerably smaller than the effect of the noise on the tone. Decreasing the noise bandwidth still further to a subcritical width reduces the asymmetry of masking somewhat, but a difference at high intensities of about 20 dB between the masking effects of an equally intense noise and tone remains. Whether the masker is a tone or noise, masking ceases when the effective energy of the masked and masking stimuli is the same.     

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.     

The fear potentiated startle effect. Blink reflex modulation as a result of classical aversive conditioning

A differential conditioning study examined whether an acoustic startle probe, presented during extinction of an aversively conditioned visual stimulus, potentiated the reflex eyeblink response in humans and whether this potentiation varied with the change in affective valence of the conditioned stimulus. Sixty college students were randomly assigned to view a series of two slides, depicting either unpleasant/highly arousing, unpleasant/moderate arousing, neutral/calm, pleasant/moderate arousing or pleasant/highly arousing scenes and objects (duration: 8 sec). During preconditioning (8 trials) and extinction (24 trials) acoustic startle probes (white noise bursts [50 ms; 95 dBA] were administered during and between slide presentation). During acquisition (16 trials) CS+ was reinforced by an electric shock. Startle response magnitudes significantly increased from preconditioning to extinction and were substantially larger to CS+. Conditioned startle reflex augmentation linearly increased with the pleasantness of the slides. Furthermore, subjects showed a greater post-conditioning increase of judged aversiveness to slides that they had previously reported to be more pleasant, exactly paralleling the startle reflex results.     

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.     

The fear potentiated startle effect

A differential conditioning study examined whether an acoustic startle probe, presented during extinction of an aversively conditioned visual stimulus, potentiated the reflex eyeblink response in humans and whether this potentiation varied with the change in affective valence of the conditioned stimulus. Sixty college students were randomly assigned to view a series of two slides, depicting either unpleasant/highly arousing, unpleasant/moderate arousing, neutral/calm, pleasant/moderate arousing or pleasant/highly arousing scenes and objects (duration: 8 sec). During preconditioning (8 trials) and extinction (24 trials) acoustic startle probes (white noise bursts [50 ms; 95 dBA] were administered during and between slide presentation). During acquisition (16 trials) CS+ was reinforced by an electric shock. Startle response magnitudes significantly increased from preconditioning to extinction and were substantially larger to CS+. Conditioned startle reflex augmentation linearly increased with the pleasantness of the slides. Furthermore, subjects showed a greater post-conditioning increase of judged aversiveness to slides that they had previously reported to be more pleasant, exactly paralleling the startle reflex results.

     

Some perceptual properties of consonants in multitalker babble

This study investigated whether consonant phonetic features or consonant acoustic properties more appropriately describe perceptual confusions among speech stimuli in multitalker babble backgrounds. Ten normal-hearing subjects identified 19 consonants, each paired with /a/, 1–19 and lui in a CV format. The stimuli were presented in quiet and in three levels of babble. Multidimensional scaling analyses of the confusion data retrieved stimulus dimensions corresponding to consonant acoustic parameters. The acoustic dimensions identified were: periodicity/burst onset, friction duration, consonant-vowel ratio, second formant transition slope, and first formant transition onset. These findings are comparable to previous reports of acoustic effects observed in white-noise conditions, and support the theory that acoustic characteristics are the relevant perceptual properties of speech in noise conditions. Perceptual effects of vowel context and level of the babble also were observed. These condition effects contrast with those previously reported for white-noise interference, and are attributed to direct masking of the low-frequency acoustic cues in the nonsense syllables by the low-frequency spectrum of the babble.     

Prepulse effects on magnitude estimation of startle-eliciting stimuli and startle responses

The present studies investigated the relationship between prepulse effects on the modification of the brainstem startle reflex and magnitude estimates of startle-eliciting stimuli. In Experiment 1, startle eyeblink responses were elicited in 24 students, half of whom were instructed to estimate the loudness of the startle stimulus (actual intensities of 80, 90, and 100 dB) and half of whom were instructed to estimate the magnitude of their eyeblink. When weak acoustic prepulses preceded the startle-eliciting stimulus, eyeblink amplitude was inhibited, and estimates of response magnitude decreased, but estimates of startle stimulus magnitude decreased only when 100-dB startle stimuli were presented. In Experiment 2, the same startle stimuli were preceded on some trials by a vibrotactile prepulse to the hand. In conditions in which startle amplitude was inhibited, startle stimulus magnitude estimates were not affected. This suggests that the effect of acoustic prepulses on 100-dB startle stimuli in Experiment 1 may have been due to loudness assimilation, an effect independent of the prepulse inhibition of startle responding.