Auditory sensitivity of an acoustic parasitoid (Emblemasoma sp., Sarcophagidae, Diptera) and the calling behavior of potential hosts.

Using field broadcasts of model male calling songs, we tested whether Tibicen pruinosa and T. chloromera (Hemiptera: Cicadidae) are candidate hosts for acoustic parasitoid flies. The model calling song of T. pruinosa attracted 90% of the flies (Sarcophagidae: Emblemasoma sp.; all larvapositing females) when broadcast simultaneously with the model T. chloromera song, a phonotactic bias reconfirmed in single song playbacks. In paired broadcasts of model T. pruinosa songs with different relative amplitudes (3 dB or 6 dB), significantly more flies were attracted to the more powerful song, a result consistent with the responses predicted by a model proposed by Forrest and Raspet [1994]. Using intracellular recordings and dye injections, we characterized the sensitivity of auditory units in sound-trapped flies. Intracellular recordings from six auditory units (5 interneurons, 1 afferent) revealed best sensitivity for frequencies near 3-4 kHz, matching the predominant spectral components of the calling songs of both species of cicada. Interestingly, although flies could be attracted to T. pruinosa broadcasts throughout the day, hourly censuses of singing males revealed that calling occurred exclusively at dusk. Furthermore, the duration of the dusk chorus in T. pruinosa was significantly shorter than the midday chorus of the less attractive song of T. chloromera. We propose that the tight temporal aggregation of the dusk chorus time could function to reduce risk from attracted parasitoids.

Hearing in mole crickets (Orthoptera: Gryllotalpidae) at sonic and ultrasonic frequencies.

We have studied auditory responses in two species of mole cricket (Scapteriscus borellii and S. abbreviatus) to determine (1) whether they show sensitivity to ultrasound, (2) whether their hearing (at both low and high frequencies) is based on the same neural circuitry as that of true crickets, and (3) whether ultrasound sensitivity in different mole cricket species varies with their ability to fly. S. borellii are sensitive to ultrasonic frequencies. There is evidence of a segregation of frequency bands in prothoracic auditory neurons. There are two pairs of omega neurons (ONs) with similar morphology to ON1 of true crickets. The two pairs of ONs differ in tuning. One pair has two sensitivity peaks: at the frequency of the calling song of this species (3 kHz), and in the ultrasonic range (25 kHz). The other pair lacks the high-frequency sensitivity and responds exclusively to frequencies in the range of the species song. These two types are not morphologically distinguishable. In S. abbreviatus, only one class of ON was found. S. abbreviatus ONs are narrowly tuned to the frequency of the species' calls. A T-neuron had the best ultrasonic frequency sensitivity in S. borellii. This cell showed a broad tuning to ultrasonic frequencies and was inhibited by low-frequency stimuli. A morphologically similar neuron was also recorded in S. abbreviatus, but lacked the high-frequency sensitivity peak of that in S. borellii. We also assessed the responses of flying S. borellii to ultrasound using field playbacks to free-flying animals. The attractiveness of broadcast calling song was diminished by the addition of an ultrasound signal, indicating that S. borellii avoid high-frequency sound. The results indicate that mole crickets process low-frequency auditory stimuli using mechanisms similar to those of true crickets. They show a negative behavioural response to high-frequency stimuli, as do true crickets, but the organization of ultrasound-sensitive auditory circuitry in mole crickets differs from that of true crickets.

A tympanal hearing organ in scarab beetles.

We describe the paired hearing organ of the scarab beetle Euetheola humilis. The auditory structures of the beetle are typical of other insect ears in that they have a thinned tympanic membrane backed by a tracheal airsac with associated chordotonal sensory structures. The tympanic membranes of the beetle are part of its cervical membrane and are located behind the head, where the cervix attaches dorsally and laterally to the pronotum. Each membrane is approximately 3 microns thick. The chordotonal sensory organ, which lies within the tracheal airsac, contains 3-8 scolopidia that attach by accessory cells directly to the tympanic membrane. Neurophysiological recordings from the neck connective of the beetle revealed that the auditory system is sensitive to frequencies between 20 and 80 kHz and has a minimum threshold of approximately 58 dB at 45 kHz. The neurophysiological audiogram is identical to the behavioral audiogram for a head roll, one behavioral component of the beetle's startle response elicited by ultrasound. Blocking experiments show that the membranous structures on the cervix are indeed the hearing organs. Neurophysiologically determined thresholds increased by more than 35 dB when drops of water covered the tympanic membranes and were essentially restored to the control level when the water was later removed. At least three other genera of Dynastinae scarabs have similar tympanum-like structures located in their cervical membranes. Behavioral and neurophysiological data show that the frequency tuning of species in two of these genera, Cyclocephala and Dyscinetus, is nearly identical to that of E. humilis. Our discovery represents only the second group of beetles known to respond to airborne sounds. However, the hearing organs of these scarab beetles differ in structure and placement from those of the tiger beetles, and thus they represent an independent evolution of auditory organs in the Coleoptera.

Ultrasound acoustic startle response in scarab beetles.

We discovered an auditory sense in a night-flying scarab beetle, Euetheola humilis, the first scarab to be shown to hear airborne sounds. In the field, beetles were captured beneath speakers broadcasting ultrasound that simulated bat echolocation pulses. Apparently, the beetles took evasive action from a potential bat predator and flew into the traps. Using another behavioral assay in laboratory studies, the beetles were sensitive to frequencies ranging from 20 to 70 kHz at levels between 60 and 70 dB SPL. One component of the behavioral response, a head roll, was graded with stimulus intensity, and the number of potentials in electromyographic recordings from muscles involved in the roll increased as stimulus intensity increased. The response latency was about 40 ms at threshold, decreasing to about 30 ms at 20 dB above threshold. The beetle's short response latency is ideally suited for predator avoidance behavior and the frequency tuning of the response suggests that it could function in evasion from insectivorous bats. The beetle's acoustic sensitivity is remarkably similar to that of other night-flying insects showing ultrasound-induced startle and it should provide these scarab beetles with a similar advance warning of predation risk.

The role of frequency selectivity in measures of auditory and vibrotactile temporal resolution.

The purpose of this study was to compare the role of frequency selectivity in measures of auditory and vibrotactile temporal resolution. In the first experiment, temporal modulation transfer functions for a sinusoidally amplitude modulated (SAM) 250-Hz carrier revealed auditory modulation thresholds significantly lower than corresponding vibrotactile modulation thresholds at SAM frequencies greater than or equal to 100 Hz. In the second experiment, auditory and vibrotactile gap detection thresholds were measured by presenting silent gaps bounded by markers of the same or different frequency. The marker frequency F1 = 250 Hz preceded the silent gap and marker frequencies after the silent gap included F2 = 250, 255, 263, 310, and 325 Hz. Auditory gap detection thresholds were lower than corresponding vibrotactile thresholds for F2 markers less than or equal to 263 Hz, but were greater than the corresponding vibrotactile gap detection thresholds for F2 markers greater than or equal to 310 Hz. When the auditory gap detection thresholds were transformed into filter attenuation values, the results were modeled well by a constant-percentage (10%) bandwidth filter centered on F1. The vibrotactile gap detection thresholds, however, were independent of marker frequency separation. In a third experiment, auditory and vibrotactile rate difference limens (RDLs) were measured for a 250-Hz carrier at SAM rates less than or equal to 100 Hz. Auditory RDLs were lower than corresponding vibrotactile RDLs for standard rates greater than 10 Hz. Combination tones may have confounded auditory performance for standard rates of 80 and 100 Hz. The results from these experiments revealed that frequency selectivity influences auditory measures of temporal resolution, but there was no evidence of frequency selectivity affecting vibrotactile temporal resolution.

Detection of silent temporal gaps in sinusoidal markers.

Gap detection thresholds were measured by forced-choice procedure for conditions where the duration of a silent gap was varied adaptively between pairs of sinusoidal markers of the same or different frequency. Frequencies of the first sinusoid in a pair of markers ranged from F1 = 500 to 4000 Hz. Second-sinusoid marker frequencies F2 included F1 = F2, and usually frequencies 2%, 5%, 24%, and 50% higher than F1. In preliminary studies the role of presentation level (E/N0) on gap detection was considered. Preliminary data revealed confounding extraneous factors arising from gating transients and from overall stimulus (i.e., markers + gap) and/or masker duration cues. In the main experiments, the contributions of these extraneous cues were evaluated with experimental designs aimed at identifying and minimizing the confounding roles of these cues in gap detection. For conditions where extraneous gating transient cues were minimized (by presenting the sinusoidal markers in a continuous noise masker with random onset phase for the second sinusoid in every pair of markers) and overall stimulus duration cues were diminished (by randomizing the duration of each marker independently), gap detection thresholds increased from 5 to 90 ms as the frequency separation between F1 and F2 was increased by half an octave. When the gap detection thresholds were treated as filter attenuation values by normalizing and converting the data into decibels, the data were closely fit by the roex filter model. On average, the listeners' performances were modeled well by a constant-percentage (7%) bandwidth filter centered on F1.

Temporal gaps in noise and sinusoids.

The ability of human observers to detect partially filled or completely silent intervals (gaps) was measured using a variety of different waveforms. The slopes of the psychometric functions for gap detection using broadband noise are dependent upon the amount of noise remaining during the gap. For completely silent intervals, the psychometric function covers a range of only 2 ms, but the psychometric functions for partially filled intervals are less steep. The detection of gaps in narrow-band noise (surrounded by complementary band-reject maskers) is strongly influenced by the signal-to-noise ratio. The signal bandwidth and center frequency also influence detectability. Gap detection improved as signal bandwidth increased, and detection improved when signal bands containing gaps were centered at higher frequencies. Detection of gaps in single components of a 21-component, equal-amplitude complex also showed lower thresholds as the frequency of the component containing the gap increased. Increasing the number of components in the complex that contained the gap improved the detectability of the gap, more so when the gaps were all presented at the same time (synchronous condition). Uncertainty about the temporal position of the gap within the observation interval made the gap more difficult to detect. This temporal uncertainty effect occurred for gaps in broadband noise, in narrow-band noise, and in sinusoidal waveforms.

Detection of partially filled gaps in noise and the temporal modulation transfer function.

Results of experiments on the detection of silent intervals, or gaps, in broadband noise are reported for normal-hearing listeners. In some preliminary experiments, a gap threshold of about 2 ms was measured. This value was independent of the duration of the noise burst, variation of the noise level on each presentation, or the temporal position of the gap within the noise burst. In the main experiments, the thresholds for partial decrements in the noise waveform as well as brief increments were determined. As predicted by a model that assumes a single fixed peak-to-valley detection ratio, thresholds for increments are slightly higher than thresholds for decrements when the signal is measured as the change in rms noise level. A first-order model describes the temporal properties of the auditory system as a low-pass filter with a 7- to 8-ms time constant. Temporal modulation transfer functions were determined for the same subjects, and the estimated temporal parameters agreed well with those estimated from the gap detection data. More detailed modeling was carried out by simulating Viemeister's three-stage temporal model. Simulations, using an initial stage bandwidth of 4000 Hz and a 3-ms time constant for the low-pass filter, generate data that are very similar to those obtained from human subjects in both modulation and gap detection.

Frequency effects in profile analysis and detecting complex spectral changes.

Seven experiments on the detectability of intensity changes in complex multitonal acoustic spectra are reported. Two general questions organize the experimental efforts. The first question is how the detectability of a change in a flat (equal energy) spectrum depends on the frequency region where a single intensive change is made. The answer is that frequency region plays a relatively minor role. Frequency changes in the midregion of the spectrum are the easiest to hear, but thresholds increase by only about 5 dB over the range from 200 to 5000 Hz. For all frequencies, the psychometric function is of the form d' = k(delta p), where k is a constant and delta p is the change in pressure. The second question is how can we predict the detectability of complex changes over the entire frequency range from the detectability of change at each separate region. Thresholds for detecting a change from a flat spectrum to a spectrum whose amplitude varies in sinusoidal ("rippled") fashion over logarithmic frequency are measured at different frequencies of ripple. The thresholds are found to be independent of ripple frequency and are 7 dB higher than predicted on the basis of an optimum combination rule.

Insect size tactics and developmental strategies.

Insect size tactics or developmental strategies are discussed in relation to decisions individuals make about when to mature. Such decisions carry with them costs and benefits in terms of that individual's reproductive success. Whenever size affects fitness, selection should act such that individuals evaluate the costs and benefits due to changes in size and should mature when the ratio of benefit to cost is maximized.Predictions about seasonal changes in adult sizes within a population are tested on two species of mole cricket, Scapteriscus acletus and vicinus. Specifically, individuals maturing in the fall should be larger than average because there is no cost associated with delayed reproduction since reproduction occurs only during spring months. Smaller than average individuals should remain in juvenile stages and get larger before reproducing. Also it is predicted that as the spring reproductive season progresses a greater proportion of smaller individuals should mature because the costs due to delaying reproduction increase. The changes in seasonal distribution of adult sizes of mole crickets support the predictions and suggest that individuals make decisions about when to mature based on costs and benefits associated with changes in size.

Profile analysis and background noise.

Spectral shape discrimination, or profile analysis, of complex waveforms (21 components) in the presence of broadband noise and special sinusoidal maskers of random amplitude was studied. The first experiment involved the discrimination between a standard flat spectrum and a "rippled" spectrum in broadband noise of different spectrum levels. Thresholds obtained under control conditions, without noise or without a standard, were used to estimate constants of an equation that predict thresholds where standard and noise are both present. The model assumes an external variance, produced by the noise, is added linearly to an internal variance caused by the flat standard. The mean squared error is less than 2 dB. The second experiment involved the detection of an increment on the center component of the 21-component standard. Added to the standard was an additional masking sinusoid of random amplitude. Both the frequency and the range of the random amplitude were varied and both showed a systematic influence on the detectability of the 1000-Hz increment.