The development of hearing sensitivity in altricial birds: the absolute thresholds of the generation of evoked potentials]

Thresholds of field L auditory evoked potentials EP were studied in 1.5-9-day-old nestlings of pied flycatcher in response to pure tone signals of different frequencies. According to the tendencies of age dynamics of auditory EP thresholds (2-9 days of life) all hearing range was divided into three separate channels: low-frequency (0.3-1.0 kHz), intermediate-(1.5-4.0 kHz) and high-frequency (5.0-8.0 kHz) ones. The widening of the hearing range in its high-frequency part was demonstrated on days 4-5 of life. The development of auditory sensitivity was shown to continue within all three channels in postembryonic period (days 2-9 post-hatching), each of the channels being characterized by its own age dynamics of auditory EP thresholds. Reproduction of tape-recorded species song during days 1-3 post-hatching resulted in significant decrease of auditory EP thresholds.     

Electrical signalling of dominance in a wild population of electric fish

Animals often use signals to communicate their dominance status and avoid the costs of combat. We investigated whether the frequency of the electric organ discharge (EOD) of the weakly electric fish, Sternarchorhynchus sp., signals the dominance status of individuals. We correlated EOD frequency with body size and found a strong positive relationship. We then performed a competition experiment in which we found that higher frequency individuals were dominant over lower frequency ones. Finally, we conducted an electrical playback experiment and found that subjects more readily approached and attacked the stimulus electrodes when they played low-frequency signals than high-frequency ones. We propose that EOD frequency communicates dominance status in this gymnotiform species.     

Fourier and electrophysiological analyses of acoustic communication in Acheta domesticus

Waveform and spectral analysis were made on the call song of the cricket, Acheta domesticus. Sounds produced by unrestrained male crickets were led directly to a computer where discrete Fourier transforms were performed on selected segments of the call song. The findings revealed essentially pure tone carrier frequencies which result from the rate at which the individual teeth of the pars stridens are struck by the plectrum. An electrophysiologically determined audiogram showed good agreement with the dominant frequency of the call song, but was less sensitive and more broadly tuned than hearing curves of most fiels crickets.     

Comparison of the physiology of the auditory receptor organs in Gryllus bimaculatus and Ephippiger ephippiger: CSD Recordings within the auditory neuropiles

The syllables of the song of the tettigoniid Ephippiger ephippiger consist of a series of short sound impulses with a broad-banded frequency spectrum. Syllables of the song of the gryllid species Gryllus bimaculatus are nearly pure tones with sharply tuned frequency maxima. A comparison of the physiology of the audiotory receptor organs of both species was carried out by using acoustical stimuli with different carrier frequencies and time-amplitude patterns. The neuronal ensemble activity of receptor cell groups of the tympanal organ was measured within the prothoracic ganglion using the CSD technique. In E. ephippiger, response maxima were found at carrier frequencies mirroring the broad frequency content of the conspecific song. The receptor cells of E. ephippiger are highly sensitive to transient sound impulses. In G. bimaculatus, the receptor cell population is more sharply tuned to the basic frequencies of the natural songs; pure tones represent more effective stimuli than transient sound signals. The causes for these speciesspecific differences are discussed with regard to probable adaptations of the receptor organs to the parameters of the conspecific songs. © 1993 John Wiley & Sons, Inc.     

Is it the Song or the Singers? Acoustic and Social Experiences Shape Adult Reproductive Tactics and Condition

When sexual signals are perceived during growth and development they can provide information regarding the social conditions likely to be encountered as an adult. Perception of cues related to the presence and density of future mates and potential competitors can result in altered adult phenotypes. Previous studies have shown that adult male Teleogryllus oceanicus field crickets from a Kauai, Hawaii population reared alone and without hearing conspecific song are more phonotactic than those reared with song. These naïve males also reduce investment in body size and immunity. Here we examined whether another source of population density information, the presence of other males, affects behavior, size, and immunity. Specifically, we examined satellite behavior as evidenced by strength of phonotaxis, body condition, and immune response in males reared singly and in groups in the presence and absence of conspecific song. Body condition did not vary with rearing density, and immune response did not vary with either acoustic environment or rearing density. Interestingly, group-housed males were more phonotactic than singly-housed males. This pattern was largely driven by the low levels of phonotaxis exhibited by males that were singly-housed in the presence of conspecific song. These findings suggest that males respond to social cues in addition to conspecific song, but that these cues do not necessarily provide concordant information.     

Rapid effects of hearing song on catecholaminergic activity in the songbird auditory pathway

Catecholaminergic (CA) neurons innervate sensory areas and affect the processing of sensory signals. For example, in birds, CA fibers innervate the auditory pathway at each level, including the midbrain, thalamus, and forebrain. We have shown previously that in female European starlings, CA activity in the auditory forebrain can be enhanced by exposure to attractive male song for one week. It is not known, however, whether hearing song can initiate that activity more rapidly. Here, we exposed estrogen-primed, female white-throated sparrows to conspecific male song and looked for evidence of rapid synthesis of catecholamines in auditory areas. In one hemisphere of the brain, we used immunohistochemistry to detect the phosphorylation of tyrosine hydroxylase (TH), a rate-limiting enzyme in the CA synthetic pathway. We found that immunoreactivity for TH phosphorylated at serine 40 increased dramatically in the auditory forebrain, but not the auditory thalamus and midbrain, after 15 min of song exposure. In the other hemisphere, we used high pressure liquid chromatography to measure catecholamines and their metabolites. We found that two dopamine metabolites, dihydroxyphenylacetic acid and homovanillic acid, increased in the auditory forebrain but not the auditory midbrain after 30 min of exposure to conspecific song. Our results are consistent with the hypothesis that exposure to a behaviorally relevant auditory stimulus rapidly induces CA activity, which may play a role in auditory responses.     

Quantitative trait loci on chromosome 5 for susceptibility to frequency-specific effects on hearing in DBA/2J mice

The DBA/2J strain is a model for early-onset, progressive hearing loss in humans, as confirmed in the present study. DBA/2J mice showed progression of hearing loss to low-frequency sounds from ultrasonic-frequency sounds and profound hearing loss at all frequencies before 7 months of age. It is known that the early-onset hearing loss of DBA/2J mice is caused by affects in the ahl (Cdh23^ahl) and ahl8 (Fscn2^ahl8) alleles of the cadherin 23 and fascin 2 genes, respectively. Although the strong contributions of the Fscn2^ahl8 allele were detected in hearing loss at 8- and 16-kHz stimuli with LOD scores of 5.02 at 8 kHz and 8.84 at 16 kHz, hearing loss effects were also demonstrated for three new quantitative trait loci (QTLs) for the intervals of 50.3–54.5, 64.6–119.9, and 119.9–137.0 Mb, respectively, on chromosome 5, with significant LOD scores of 2.80–3.91 for specific high-frequency hearing loss at 16 kHz by quantitative trait loci linkage mapping using a (DBA/2J × C57BL/6J) F₁ × DBA/2J backcross mice. Moreover, we showed that the contribution of Fscn2^ahl8 to early-onset hearing loss with 32-kHz stimuli is extremely low and raised the possibility of effects from the Cdh23^ahl allele and another dominant quantitative trait locus (loci) for hearing loss at this ultrasonic frequency. Therefore, our results suggested that frequency-specific QTLs control early-onset hearing loss in DBA/2J mice.     

Ultrasonic signalling by a Bornean frog

Among anuran amphibians, only two species, Odorrana tormota and Huia cavitympanum, are known to possess recessed tympanic membranes. Odorrana tormota is the first non-mammalian vertebrate demonstrated to communicate with ultrasonic frequencies (above 20 kHz), and the frogs' sunken tympana are hypothesized to play a key role in their high-frequency hearing sensitivity. Here we present the first data on the vocalizations of H. cavitympanum. We found that this species emits extraordinarily high-frequency calls, a portion of which are comprised entirely of ultrasound. This represents the first documentation of an anuran species producing purely ultrasonic signals. In addition, the vocal repertoire of H. cavitympanum is highly variable in frequency modulation pattern and spectral composition. The frogs' use of vocal signals with a wide range of dominant frequencies may be a strategy to maximize acoustic energy transmission to both nearby and distant receivers. The convergence of these species' call characteristics should stimulate additional, phylogenetically based studies of other lower vertebrates to provide new insight into the mechanistic and evolutionary foundations of high-frequency hearing in all vertebrate forms.     

Encoding properties of auditory neurons in the brain of a soniferous damselfish: response to simple tones and complex conspecific signals

The fish auditory system encodes important acoustic stimuli used in social communication, but few studies have examined response properties of central auditory neurons to natural signals. We determined the features and responses of single hindbrain and midbrain auditory neurons to tone bursts and playbacks of conspecific sounds in the soniferous damselfish, Abudefduf abdominalis. Most auditory neurons were either silent or had slow irregular resting discharge rates <20 spikes s⁻¹. Average best frequency for neurons to tone stimuli was ~130 Hz but ranged from 80 to 400 Hz with strong phase-locking. This low-frequency sensitivity matches the frequency band of natural sounds. Auditory neurons were also modulated by playbacks of conspecific sounds with thresholds similar to 100 Hz tones, but these thresholds were lower than that of tones at other test frequencies. Thresholds of neurons to natural sounds were lower in the midbrain than the hindbrain. This is the first study to compare response properties of auditory neurons to both simple tones and complex stimuli in the brain of a recently derived soniferous perciform that lacks accessory auditory structures. These data demonstrate that the auditory fish brain is most sensitive to the frequency and temporal components of natural pulsed sounds that provide important signals for conspecific communication.     

Useless Hearing in Male Emblemasoma auditrix (Diptera,Sarcophagidae) – A Case of Intralocus Sexual Conflict during Evolution of a Complex Sense Organ?

Sensory modalities typically are important for both sexes, although sex-specific functional adaptations may occur frequently. This is true for hearing as well. Consequently, distinct behavioural functions were identified for the different insect hearing systems. Here we describe a first case, where a trait of an evolutionary novelty and a highly specialized hearing organ is adaptive in only one sex. The main function of hearing of the parasitoid fly Emblemasoma auditrix is to locate the host, males of the cicada species Okanagana rimosa, by their calling song. This task is performed by female flies, which deposit larvae into the host. We show that male E. auditrix possess a hearing sense as well. The morphology of the tympanal organ of male E. auditrix is rather similar to the female ear, which is 8% broader than the male ear. In both sexes the physiological hearing threshold is tuned to 5 kHz. Behavioural tests show that males are able to orient towards the host calling song, although phonotaxis often is incomplete. However, despite extensive observations in the field and substantial knowledge of the biology of E. auditrix, no potentially adaptive function of the male auditory sense has been identified. This unique hearing system might represent an intralocus sexual conflict, as the complex sense organ and the behavioural relevant neuronal network is adaptive for only one sex. The correlated evolution of the sense organ in both sexes might impose substantial constraints on the sensory properties of the ear. Similar constraints, although hidden, might also apply to other sensory systems in which behavioural functions differ between sexes.     

Cricket phonotaxis: localization depends on recognition of the calling song pattern

Summary Calling song with a carrier frequency of 5 kHz evokes positive phonotaxis in female crickets,Gryllus bimaculatus, when presented at an azimuth. In contrast, a continuous tone of 4.7 kHz in the same position when paired with calling song from above leads to negative phonotaxis. Under open-loop conditions, when a tethered animal runs on a paired tread wheel, characteristic curves are produced with the stable equilibrium point towards or away from the stimulus, respectively (Fig. 3).In order to understand this sign reversal at the neuronal level, directional characteristics of the ascending acoustic inter neurons AN1 and AN2 were measured using extracellular recordings from the cervical connectives.Taking the mean spike rate of the interneurons as a measure for their excitation, the function relating response magnitude to stimulus direction for calling song corresponds well to the behavioural characteristic curve (Fig. 5). The response function obtained using a continuous tone with simultaneous presentation of calling song from above is similar (Fig. 5) and hence does not correspond to the inverse behavioural characteristic curve.However, consideration of the extent to which the temporal parameters of the calling song (syllables and chirps) are reflected in the neuronal response (amplitudes of the Fourier components) leads to characteristic curves for AN1 and AN2 which are in good agreement with the behaviour for stimulation with calling song as well as for the continuous tone experiment (Fig. 8). In addition, the neural response curves correspond to the behaviour in showing smaller amplitudes when a continuous tone rather than the calling song is presented on the horizon (Fig. 8).From these data we conclude that the activity in interneurons AN1 and AN2 does not directly guide orientation in mating behaviour but first is filtered by a mechanism tuned to the frequency of syllables and/or chirps. According to this hypothesis recognition of conspecific song and localization proceed sequentially inGryllus.     

Lateralization of Travelling Wave Response in the Hearing Organ of Bushcrickets

Travelling waves are the physical basis of frequency discrimination in many vertebrate and invertebrate taxa, including mammals, birds, and some insects. In bushcrickets (Tettigoniidae), the crista acustica is the hearing organ that has been shown to use sound-induced travelling waves. Up to now, data on mechanical characteristics of sound-induced travelling waves were only available along the longitudinal (proximal-distal) direction. In this study, we use laser Doppler vibrometry to investigate in-vivo radial (anterior-posterior) features of travelling waves in the tropical bushcricket Mecopoda elongata. Our results demonstrate that the maximum of sound-induced travelling wave amplitude response is always shifted towards the anterior part of the crista acustica. This lateralization of the travelling wave response induces a tilt in the motion of the crista acustica, which presumably optimizes sensory transduction by exerting a shear motion on the sensory cilia in this hearing organ.     

Song discrimination by nestling collared flycatchers during early development

Pre-zygotic isolation is often maintained by species-specific signals and preferences. However, in species where signals are learnt, as in songbirds, learning errors can lead to costly hybridization. Song discrimination expressed during early developmental stages may ensure selective learning later in life but can be difficult to demonstrate before behavioural responses are obvious. Here, we use a novel method, measuring changes in metabolic rate, to detect song perception and discrimination in collared flycatcher embryos and nestlings. We found that nestlings as early as 7 days old respond to song with increased metabolic rate, and, by 9 days old, have increased metabolic rate when listening to conspecific when compared with heterospecific song. This early discrimination between songs probably leads to fewer heterospecific matings, and thus higher fitness of collared flycatchers living in sympatry with closely related species.     

Hearing conspecific vocal signals alters peripheral auditory sensitivity

We investigated whether hearing advertisement calls over several nights, as happens in natural frog choruses, modified the responses of the peripheral auditory system in the green treefrog, Hyla cinerea. Using auditory evoked potentials (AEP), we found that exposure to 10 nights of a simulated male chorus lowered auditory thresholds in males and females, while exposure to random tones had no effect in males, but did result in lower thresholds in females. The threshold change was larger at the lower frequencies stimulating the amphibian papilla than at higher frequencies stimulating the basilar papilla. Suprathreshold responses to tonal stimuli were assessed for two peaks in the AEP recordings. For the peak P1 (assessed for 0.8–1.25 kHz), peak amplitude increased following chorus exposure. For peak P2 (assessed for 2–4 kHz), peak amplitude decreased at frequencies between 2.5 and 4.0 kHz, but remained unaltered at 2.0 kHz. Our results show for the first time, to our knowledge, that hearing dynamic social stimuli, like frog choruses, can alter the responses of the auditory periphery in a way that could enhance the detection of and response to conspecific acoustic communication signals.     

Ultrasonic singing by the blue-throated hummingbird: a comparison between production and perception

Blue-throated hummingbirds produce elaborate songs extending into the ultrasonic frequency range, up to 30 kHz. Ultrasonic song elements include harmonics and extensions of audible notes, non-harmonic components of audible syllables, and sounds produced at frequencies above 20 kHz without corresponding hearing range sound. To determine whether ultrasonic song elements function in intraspecific communication, we tested the hearing range of male and female blue-throated hummingbirds. We measured auditory thresholds for tone pips ranging from 1 kHz to 50 kHz using auditory brainstem responses. Neither male nor female blue-throated hummingbirds appear to be able to hear above 7 kHz. No auditory brainstem responses could be detected between 8 and 50 kHz at 90 dB. This high-frequency cutoff is well within the range reported for other species of birds. These results suggest that high-frequency song elements are not used in intraspecific communication. We propose that the restricted hummingbird hearing range may exemplify a phylogenetic constraint.     

Males of <Emphasis Type="Italic">Hypsiboas goianus</Emphasis> (Anura; Hylidae) do not assess neighbor fighting ability through acoustic interactions

In frogs, acoustic signals are the most important communication mechanism, since they may be used in several social contexts. In many anurans, the dominant frequency of calls is negatively related to body size, and in such species, this spectral parameter may be considered a good predictor of fighting ability. We experimentally investigated the vocal behavior of 30 male Hypsiboas goianus in central Brazil to answer the following questions: (1) Do males change the acoustic parameters of their calls in response to conspecific intruders? and (2) Does the acoustic behavior of H. goianus depend on the simulated body size of their opponent? We used playback of synthesized calls with high (3573 Hz) and low (3123 Hz) dominant frequency to simulate small and large males, respectively. Males reduced the rate of advertisement calling in response to playback but did not change vocal behavior in response to low-frequency and high-frequency playback. So, while males adjust their calling activity in response to simulated conspecifics, there was no evidence that they assess the fighting ability of their opponents through acoustic interactions.     

Species Recognition During Substrate-Borne Communication in <Emphasis Type="Italic">Nezara viridula</Emphasis> (L.) (Pentatomidae: Heteroptera)

The information code in the temporal and spectral characteristics of the substrate-borne communication signals produced by insects has been primarily studied in insects in the suborder Auchenorrhyncha. In the present study we investigated which of the female calling song (FCS) parameters in Nezara viridula (L.) (Heteroptera, Pentatomidae) are essential for recognition by conspecific males. In playback experiments we measured male vibrational responsiveness to FCS signals varying in the durations of pulse trains and inter-pulse train intervals, repetition times, duty cycles, and dominant frequencies, and determined the preference range for each specific parameter. Males were able to distinguish songs of different temporal and frequency parameters and responded best to values characteristic of the song of conspecific females. Signal recognition is achieved on the basis of two temporal filters tuned to the durations of the pulse train and inter-pulse train interval. Males responded best to the dominant frequency characteristic of conspecific songs, which are tuned to the resonant properties of the herbaceous plants used for intraspecific signal transmission during communication.     

The acoustic behaviour of the bushcricket Tettigonia cantans II. Transmission of airborne-sound and vibration signals in the biotope

The airborne-sound and the vibratory signals produced by stridulating $\text{Tettigonia cantans}$ males, and the transmission of these signals in the natural biotope were investigated.The song of $\text{T. cantans}$ is composed of repeated uniform syllables with a rate of ca. 30/sec. Intensity approaches 100 dB SPL, 10 cm away from the animal. The spectrum shows three dominant frequency ranges around 8, 16 and 32 kHz.Airborne transmission of the song in such vegetation layers as are found in the biotopes of $\text{T. cantans}$ shows an excess attenuation which increases with frequency. The relative intensities of the frequency components of the song vary as a result of the kind of vegetation, the positions of emitter and receiver, and the separation distance. These relative differences in intensity may be useful during the phonotactic approach to conspecific partners, providing a measure of the distance from the sound source.Stridulating males also produce vibratory signals in the plants they sit on. The spectrum of these signals includes frequencies up to 8 kHz, the first dominant frequency of the song: low frequency components are induced in the plants via the legs and abdomen of the animal. The vibratory signals are transmitted mainly in the form of bending waves. Near the animal, amplitude modulation corresponds to that of the song. At greater distances, reflections and frequency-dependent propagation velocities, cause distortions of this time pattern. Transmission depends greatly on the mechanical properties of the particular plant, attenuation values of 20–50 dB/m being found. Nevertheless, in most cases, vibratory signals may be perceived up to 1.5 – 2 metres away from a stridulating male.