The modulation rate transfer function of a harbour porpoise (Phocoena phocoena)

During echolocation, toothed whales produce ultrasonic clicks at extremely rapid rates and listen for the returning echoes. The auditory brainstem response (ABR) duration was evaluated in terms of latency between single peaks: 5.5 ms (from peak I to VII), 3.4 ms (I–VI), and 1.4 ms (II–IV). In comparison to the killer whale and the bottlenose dolphin, the ABR of the harbour porpoise has shorter intervals between the peaks and consequently a shorter ABR duration. This indicates that the ABR duration and peak latencies are possibly related to the relative size of the auditory structures of the central nervous system and thus to the animal’s size. The ABR to a sinusoidal amplitude modulated stimulus at 125 kHz (sensitivity threshold 63 dB re 1 μPa rms) was evaluated to determine the modulation rate transfer function of a harbour porpoise. The ABR showed distinct envelope following responses up to a modulation rate of 1,900 Hz. The corresponding calculated equivalent rectangular duration of 263 μs indicates a good temporal resolution in the harbour porpoise auditory system similar to the one for the bottlenose dolphin. The results explain how the harbour porpoise can follow clicks and echoes during echolocation with very short inter click intervals.     

The effect of click rate on latency and interpeak interval of the brain-stem auditory evoked potentials in children from birth to 6 years

Latency and interpeak interval of the brain-stem auditory evoked potentials at different click rates were measured in 80 healthy children from birth to 6 years, and 21 adults. Clicks were presented at 10, 30, 50, 70 and 90/sec, and 70, 40 and 20 db HL. At high stimulus intensity (70 dB SL), all latencies of waves I, III and V and the I–V, I–III and III–V intervals showed a progressive prolongation with increasing repetition rate. The latency- and the interval-rate functions were similar for all age groups but their slopes were slightly steeper in younger than in older. As click rate increased from 10/sec to 90/sec, the latencies of waves I, III and V at different age groups were prolonged by 4–10%, 9–13% and 12–15% respectively, and the intervals of I–V, I–III and III–V were prolonged by 15–16%, 8–16% and 14–24% respectively. The mean increments of wave V latency and I–V interval in different age groups were 0.404–0.575 and 0.332–0.526 msec respectively with increasing click rate from 10 to 50/sec, and 0.697–1.009 and 0.629–0.776 msec respectively with increasing click rate from 10 to 90/sec. The younger the age the larger the absolute increments for all these BAEP parameters, but the increasing rates for a BAEP measure were similar among different age groups, exhibiting no age-dependent differences. The III–V/I–III interval ration in most age groups was increased by 3–10% with increasing click rate from 10 to 90/sec, suggesting that the III–V interval was affected by stimulus rate slightly more than I–III interval.At moderate (40 dB HL) and low (20 dB SL) intensity, all waves and intervals showed similar latency- and interval-rate functions to those at high intensity. This demonstrates that the shifting latencies and interpeak intervals with increasing click rate appeared to be independent of the stimulus intensities.     

Whole-head mapping of middle-latency auditory evoked magnetic fields

We recorded middle-latency auditory evoked magnetic fields from 9 healthy subjects with a 122-channel whole-head SQUID gradiometer. The stimuli were click triplets, 2.5 msec in total duration, delivered alternately to the two ears once every 333 msec. Contralateral clicks elicited P30m responses in 16 and P50m responses in 12 out of 18 hemispheres studied; ipsilateral clicks did so in 7 and 13 hemispheres, respectively. The field patterns were satisfactorily explained by current dipoles in 16 and 4 hemispheres for contra- and ipsilateral P30m, and in 4 and 10 hemispheres for contra- and ipsilateral P50m. The peak latencies of P30m and P50m were not affected by stimulation side. The results show that middle-latency auditory evoked responses receive a strong contribution from auditory cortical structures, and that differences of input latency to cortical auditory areas, evaluated from MLAEF latencies, do not explain the latency differences seen in late auditory evoked fields to contralateral vs. ipsilateral stimulation.     

Two measures of temporal resolution in brown-headed cowbirds (<Emphasis Type="Italic">Molothrus ater</Emphasis>)

Studies of auditory temporal resolution in birds have traditionally examined processing capabilities by assessing behavioral discrimination of sounds varying in temporal structure. Here, temporal resolution of the brown-headed cowbird (Molothrus ater) was measured using two auditory evoked potential (AEP)-based methods: auditory brainstem responses (ABRs) to paired clicks and envelope following responses (EFRs) to amplitude-modulated tones. The basic patterns observed in cowbirds were similar to those found in other songbird species, suggesting similar temporal processing capabilities. The amplitude of the ABR to the second click was less than that of the first click at inter-click intervals less than 10 ms, and decreased to 30% at an interval of 1 ms. EFR amplitude was generally greatest at modulation frequencies from 335 to 635 Hz and decreased at higher and lower modulation frequencies. Compared to data from terrestrial mammals these results support recent behavioral findings of enhanced temporal resolution in birds. General agreement between these AEP results and behaviorally based studies suggests that AEPs can provide a useful assessment of temporal resolution in wild bird species.     

Temporal resolution of the Risso’s dolphin, Grampus griseus, auditory system

Toothed whales and dolphins (Odontocetes) are known to echolocate, producing short, broadband clicks and receiving the corresponding echoes, at extremely rapid rates. Auditory evoked potentials (AEP) and broadband click stimuli were used to determine the modulation rate transfer function (MRTF) of a neonate Risso’s dolphin, Grampus griseus, thus estimating the dolphin’s temporal resolution, and quantifying its physiological delay to sound stimuli. The Risso’s dolphin followed sound stimuli up to 1,000 Hz with a second peak response at 500 Hz. A weighted MRTF reflected that the animal followed a broad range of rates from 100 to 1,000 Hz, but beyond 1,250 Hz the animal’s hearing response was simply an onset/offset response. Similar to other mammals, the dolphin’s AEP response to a single stimulus was a series of waves. The delay of the first wave, PI, was 2.76 ms and the duration of the multi-peaked response was 4.13 ms. The MRTF was similar in shape to other marine mammals except that the response delay was among the fastest measured. Results predicted that the Risso’s dolphin should have the ability to follow clicks and echoes while foraging at close range.     

白噪声对听觉脑干电反应(ABR)和耳蜗电图(ECochG)的影响

本文通过20例听力正常人和10例听力正常豚鼠研究了白噪声对耳蜗电图(ECochG)和听觉脑干电反应(ABR)的干涉作用。实验结果表明,白噪声比短声(信号)的声强级低30dB(SL)以上时,ECochG和ABR的振幅仅轻微减小。白噪声与短声的声强级相等时,ECochG与ABR的振幅和出现率会明显受到干涉而减小,甚至完全消失。但是,此时的耳蜗微音器电位(CM)并未观察到有明显的变化。这意味着白噪声对ECochG和ABR的干涉作用主要与围绕毛细胞基底部的突触产生的抑制密切相关。由于白噪声对ABR各波的干涉有些差异,所以认为这种抑制,可能既包括脑中抑制也包括侧方抑制。     

Auditory brain stem responses in characterization of dolphin hearing

Summary Auditory brain stem responses (ABR) were recorded from the head surface of non-anesthetized and non-relaxed bottle-nosed dolphins, Tursiops truncatus. The region of best ABR recording was shown to be located 6–9 cm caudal to the blowhole. The threshold values were about 1 mPa for noise bursts and –3 dB re 1 mPa for tone bursts of the optimal frequency (80 kHz). The maximum frequency at which ABR could be evoked was 140 kHz. The duration of temporal summation reached 0.5 ms at intensities near the threshold and decreased with an increase in intensity. When the stimuli were paired clicks of the same intensity, the time to complete recovery from the second response was about 5 ms, while that to its 50% recovery was 0.7 ms. When the conditioning click exceeded the testing one in intensity, prolongation of the recovery period was observed. A 40-dB intensity difference led to an approximately 10-fold prolongation of this period.Abbreviations ABR auditory brain stem response - EP evoked potential     

论听觉诱发电位快成分中瞬态反应与稳态反应的相互关系

实验和计算表明,由短声串所引起的头皮诱发电位快成分中,前面的短声对后面短声所诱发的反应有掩蔽作用;短声串所诱发的整个反应,就是每个短声在它之前短声的掩蔽作用下所引起的反应之和.这一结论,看来也可推广到频率跟随反应以至更一般的听觉诱发电位快成分的情形中去.     

Auditory evoked responses to rhythmic sound pulses in dolphins

The ability of auditory evoked potentials to follow sound pulse (click or pip) rate was studied in bottlenosed dolphins. Sound pulses were presented in 20-ms rhythmic trains separated by 80-ms pauses. Rhythmic click or pip trains evoked a quasi-sustained response consisting of a sequence of auditory brainstem responses. This was designated as the rate-following response. Rate following response peak-to-peak amplitude dependence on sound pulse rate was almost flat up to 200 s⁻¹, then displayed a few peaks and valleys superimposed on a low-pass filtering function with a cut-off frequency of 1700 s⁻¹ at a 0.1-amplitude level. Peaks and valleys of the function corresponded to the pattern of the single auditory brain stem response spectrum; the low-pass cut-off frequency was below the auditory brain stem response spectrum bandwidth. Rate-following response frequency composition (magnitudes of the fundamental and harmonics) corresponded to the auditory brain stem response frequency spectrum except for lower fundamental magnitudes at frequencies above 1700 Hz. These regularities were similar for both click and pip trains. The rate-following response to steady-state rhythmic stimulation was similar to the rate-following response evoked by short trains except for a slight amplitude decrease with the rate increase above 10 s⁻¹. The latter effect is attributed to a long-term rate-dependent adaptation in conditions of the steady-state pulse stimulation. Accepted: 18 June 1998     

CHARACTERISTICS OF MINKE WHALE (BALAENOPTERA ACUTOROSTRATA) PULSE TRAINS RECORDED NEAR PUERTO RICO

A nine-day acoustic and visual survey was conducted off the West Indies in March 1994 to study the pulse trains that were detected on SOSUS arrays throughout winter in deep water between the West Indies and Bermuda. During the survey, pulse train sounds were consistently recorded in an area 190–350 km northeast of Puerto Rico. Vocalizing animals were never visually observed, but visual sighting conditions were often poor and observation height was low. Pulse trains occurred in two basic forms. The "speed-up" pulse train was characterized by an accelerating series of pulses with energy in the 200–400 Hz band, with individual pulses lasting 40-60 msec. Speedup pulse trains started with average pulse rates of 1.5 pulses/sec, lasted 43.7 ± 6.0 sec, and ended with average pulse rates of 2.8 pulses/sec. The less common "slow-down" pulse train was characterized by a decelerating series of pulses with energy in the 250-350 Hz band. Slow-down pulse trains started at pulse rates averaging 4.5 pulses/sec, lasted 60.9 ± 5.8 sec, and ended with average pulse rates of 2.9 pulses/sec. We believe the recorded pulse trains are from minke whales based on careful reanalysis of, and comparison to, minke whale pulse-train sounds recorded in the Caribbean by Winn and Perkins (1976).     

THE CLICK-SOUNDS OF NARWHALS (MONODON MONOCEROS) IN INGLEFIELD BAY, NORTHWEST GREENLAND

We studied the sounds of narwhals ( Monodon monoceros ) foraging in the open waters in Northwest Greenland. We used a linear, vertical array of three hydrophones (depth 10 m, 30 m, 100 m) with a fourth hydrophone (depth 30 m) about 20 m from the vertical array. A smaller fifth hydrophone (depth 2 m) allowed for registering frequencies up to 125 kHz (± 2 dB) when signals were recorded at 762 mm/set on an instrumentation tape recorder. Clicks were the prevalent signals, but we heard whistles occasionally. We separated the clicks into two classes: click trains that had rates of 3-10 clicks/sec and click bursts having rates of 110-150 clicks/sec. The spectra of train clicks had maximum amplitudes at 48 ± 10 kHz and a duration of 29 ± 6 psec. The spectra of burst clicks had maximum amplitudes at 19 ± 1 kHz and a duration of 40 ± 3 psec. By analogy with other dolphin species, narwhals presumably use the clicks for echolocation during orientation and for locating prey. The narwhal click patterns resemble those of insectivorous bats. Click trains might correspond to bat searching signals and click bursts to the bat's terminal "buzz", emitted just before prey capture.     

Responses of motor cortex neurons of the cat to auditory stimulation and their role in the instrumental food reflex

The responses of motor cortex neurons in the cat to the presentation of a single auditory click and a series of 10 clicks presented with 1,000/sec frequency were studied under conditions of chronic experiments before and after the development of an instrumental food reflex. After reflex development a single presentation of a positive conditioned stimulus (single click) markedly influenced for 7 sec the appearance of instrumental movements. At the same time, the immediate responses of motor cortex neurons to presentation of the conditioned auditory stimulus had no impact on the appearance in the motor cortex of discharges leading to the realization of instrumental movements. Consequently, motor cortex neurons do not require activation from afferent sensory inputs for the generation of such discharges. The immediate neuronal responses to conditioned stimulation did not inhibit the realization of the instrumental reflex. It is proposed that they are associated with the realization of motor function in the unconditioned defensive response evoked by the presentation of an auditory stimulus. The presence or absence of responses to auditory conditioned stimulation was dependent upon the signal meaning of the stimulus, its physical parameters, and the degree of excitability of the animal.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 4, pp. 539–550, July–August, 1985.     

Comparison of evoked potentials to tones and clicks recorded simulataneously by multiple electrodes from the auditory cortex in unanesthetized cats

Evoked potentials to tones and clicks were recorded simultaneously from seven points of the auditory cortex and one or two points of the somatosensory cortex in unanesthetized cats. Comparison of evoked potentials to tones of equal loudness in the 250–7000 Hz band showed no common pattern of cortical tonotopic distribution. However, an individual dependence of the components of the evoked potential on pitch and on localization of the recording point exists for each animal. With a change in stimulus intensity the absolute and relative values of these components of the evoked potential vary. The initial positive waves are the most variable; besides the two waves already known a third, intermediate wave, particulary sensitive to loudness, was discovered. The negative wave of the primary response increases proportionally to loudness. Evoked potentials to clicks are more uniform over the auditory cortex and more stable than those to tones. Responses appeared in the somatosensory cortex to loud stimuli, more regularly to clicks than to tones. It is concluded that the parameter of pitch is reflected in the cat cortex as a complex spatially-individual distribution of the amplitude and time parameters of the evoked potentials.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 7, No. 2, pp. 115–125, March–April, 1975.     

The auditory P50 response is normal in Alzheimer's disease when measured via a paired click paradigm

Recent findings of missing or markedly attenuated P50 (or P1) auditory ERPs in Alzheimer's disease (AD) patients suggest this may be a useful diagnostic and/or prognostic marker of AD cholinergic deficits. Those studies used repetitive 1/sec clicks. Given P50's long recovery time, all but the first click in that paradigm was presented during the recovery of the P50 generation system from the response to the prior click. We studied 8 AD patients and 17 elderly controls using a paradigm incorporating 7–8 sec intervals between clicks, which allows examination of P50 generation separate from P50 recovery. With the long inter-click interval, we identified P50 responses in most AD patients and controls, and found no difference in P50 amplitude between groups. These results suggest that if there is a P50 deficit in AD patients, it is the result of the accumulative effect of repetitive stimulation, rather than a primary deficit in P50 generation.     

High-rate sequential sampling of auditory brain-stem and somatosensory evoked responses in hypoxia

We developed a high-rate sequential recording technique that allowed simultaneous measurements of both auditory brain-stem response (ABR) and somatosensory evoked potential (SEP) every 10 sec. Using this method, a transient increase in amplitude of all the ABR and SEP components in response to hypoxia in dogs could be detected. The increase in amplitude preceded the prolongation of latency. Our study showed that there were succesive changes of evoked potentials in response to hypoxia. A transient increase in amplitude is the first to occur, followed by a latency prolongation and an amplitude decrease for both ABRs and SEPs.     

Responses of auditory cortical neurons to paired clicks

In cats immobilized with tubocurarine, a paired-click method was used to determine the duration of the refractory period of 75 auditory cortical neurons responding to clicks with a latent period of up to 30 msec. Sixty-eight of the neurons exhibited no spontaneous activity, while in the other seven spontaneous activity was infrequent and irregular. It was found that a click makes responding neurons refractory to a second click for a long time. The duration of this refractory period is 3 to 700 msec; it is constant for each neuron, but varies from one neuron to another. A direct relationship was found between the number of neurons responding to the second click and the interval between the first and second clicks: the shorter the interval the fewer neurons respond to the second click. It is postulated that this dependence lies at the basis of the neurophysiological mechanism of perception and discrimination of short time intervals.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 2, No. 3, pp. 227–235, May–June, 1970.     

AEPs at the onset and offset of repetitive sound modulation,due to mismatch with the contents of an auditory sensory store

Long latency auditory evoked potentials (AEPs), chiefly consisting of a negative peak at about 150 msec and a positivity at 250 msec, were recorded at the beginning and end of periods during which the interaural time difference of binaural noise was switched between 0.0 and 0.8 msec at a fast rate (ISI = 50 or 25 msec) or the frequency of continuous binaural clicks was switched between 167 and 200 Hz every 80, 50 or 25 msec. In the latter case the offset responses occurred later than onset by a mean of 89, 47 and 27 msec respectively, suggesting they were probably generated at the moment the next switch was expected but failed to occur.The offset responses must be non-specific with respect to the interaural delay or the frequency of clicks, since neurones which respond to particular delays or frequencies and are made refractory by a rapid rate of stimulation should not suddenly become less so at the last in a series of identical stimuli, or be activated by the absence of a further event. It is proposed that the potentials are due to a higher order of neurone which automatically responds to the occurrence of a “mismatch” between the immediate sound and an image of that which was previously present, encoded in a short-term sensory store. In addition to frequency content and interaural delay, the image must contain information about the temporal modulation pattern of the sound over the previous few seconds.     

Dynamics of potentials evoked in the auditory pathway and reticular formation of the cat. Studies during waking and sleep stages

Averaged evoked potentials in the inferior colliculus (IC), medial geniculate nucleus (MG) and reticular formation (RF) of chronically implanted and freely moving cats were measured using auditory step functions in the form of tone bursts of 2000 Hz. The most prominent components of the AEP of the inferior colliculus were a positive wave of 13 msec and a negative wave of 40–55 msec latency. The AEP of the medial geniculate nucleus was characterized by a large negative wave peaking at 35–40 msec. During spindle sleep and slow wave sleep stages changes in the AEPs of both nuclei occured.Transient evoked responses of the inferior colliculus, medial geniculate nucleus and reticular formation were transformed to the frequency domain using the Laplace transform (one sided Fourier transform) in order to obtain frequency characteristics of the systems under study. The amplitude characteristics of IC, MG. and RF obtained in this way revealed maxima in alpha (8–13 Hz), beta (18–35 Hz) and higher frequency (50–80 Hz) ranges. During spindle sleep stage a maximum in the theta frequency range (3–8 Hz) and during slow wave sleep maximum in the delta (1–3 Hz) frequency range appeared in the amplitude characteristics of these nuclei.The amplitude characteristics of the inferior colliculus and medial geniculate nucleus were compared with the amplitude characteristics of other brain structures. The comparison of AEPs and amplitude frequency characteristics obtained using these AEPs reveals that the existence of a number of peaks (waves) with different latencies in the time course does not necessarily indicate the existence of different functional structures or neural groups giving rise to these waves. The entire time course of evoked potentials and not the number and latencies of the waves, carries, the whole information concerning different activities and frequency selectivities of brain structures.Supported by Turkish Scientific and Technical Research Council Grant TAG-266.Presented in Part at the VIIIth International Congress of Electroencephalography and Clinical Neurophysiology in Marseilles, September 1–7, 1973.     

Auditory temporal resolution of a wild white-beaked dolphin (<Emphasis Type="Italic">Lagenorhynchus albirostris</Emphasis>)

Adequate temporal resolution is required across taxa to properly utilize amplitude modulated acoustic signals. Among mammals, odontocete marine mammals are considered to have relatively high temporal resolution, which is a selective advantage when processing fast traveling underwater sound. However, multiple methods used to estimate auditory temporal resolution have left comparisons among odontocetes and other mammals somewhat vague. Here we present the estimated auditory temporal resolution of an adult male white-beaked dolphin, (Lagenorhynchus albirostris), using auditory evoked potentials and click stimuli. Ours is the first of such studies performed on a wild dolphin in a capture-and-release scenario. The white-beaked dolphin followed rhythmic clicks up to a rate of approximately 1,125–1,250 Hz, after which the modulation rate transfer function (MRTF) cut-off steeply. However, 10% of the maximum response was still found at 1,450 Hz indicating high temporal resolution. The MRTF was similar in shape and bandwidth to that of other odontocetes. The estimated maximal temporal resolution of white-beaked dolphins and other odontocetes was approximately twice that of pinnipeds and manatees, and more than ten-times faster than humans and gerbils. The exceptionally high temporal resolution abilities of odontocetes are likely due primarily to echolocation capabilities that require rapid processing of acoustic cues.