Independence of Evoked Vocal Responses from Stimulus Direction in Burrowing Frogs Eupsophus (Leptodactylidae)

Localization of a sound source is important for animals in mating contexts: females generally orient towards signalling males, and males can estimate the position and quality of potential rivals. In anurans, the effect of sound direction on evoked vocal responses has been studied in males of Rana catesbeiana, which alter their vocal responses depending on the location of the stimulus. The current study explored the effects of sound direction in Eupsophus calcaratus, a frog that calls from inside burrows having resonance that would hinder the localization of incoming sounds. The vocal responses of 11 males to synthetic imitations of the conspecific advertisement call broadcast from loudspeakers positioned in front, to the right and left from the burrow openings were similar in terms of call rate, duration and latency. The invariance of the vocal responses indicates that for burrowing male frogs engaged in chorusing behaviour, the specific location of an opponent does not alter the persistence of vocal activity.     

Passive sound localization of prey by the pallid bat (Antrozous p. pallidus)

Summary The pallid bat (Antrozous p. pallidus) uses passive sound localization to capture terrestrial prey. This study of captive pallid bats examined the roles of echolocation and passive sound localization in prey capture, and focused on their spectral requirements for accurate passive sound localization.Crickets were used as prey throughout these studies. All tests were conducted in dim, red light in an effort to preclude the use of vision. Hunting performance did not differ significantly in red light and total darkness, nor did it differ when visual contrast between the terrestrial prey and the substrate was varied, demonstrating that the bats did not use vision to locate prey.Our bats apparently used echolocation for general orientation, but not to locate prey. They did not increase their pulse emission rate prior to prey capture, suggesting that they were not actively scanning prey. Instead, they required prey-generated sounds for localization. The bats attended to the sound of walking crickets for localization, and also attacked small, inanimate objects dragged across the floor. Stationary and/or anesthetized crickets were ignored, as were crickets walking on substrates that greatly attenuated walking sounds. Cricket communication sounds were not used in prey localization; the bats never captured stationary, calling crickets.The accuracy of their passive sound localization was tested with an open-loop passive sound localization task that required them to land upon an anesthetized cricket tossed on the floor. The impact of a cricket produced a single 10–20 ms duration sound, yet with this information, the bats were able to land within 7.6 cm of the cricket from a maximum distance of 4.9 m. This performance suggests a sound localization accuracy of approximately ±1° in the horizontal and vertical dimensions of auditory space. The lower frequency limit for accurate sound localization was between 3–8 kHz. A physiological survey of frequency representation in the pallid bat inferior colliculus suggests that this lower frequency limit is around 5 kHz.     

Phonotactic behaviour and vertical sound source localisation of the parasitoid fly Emblemasoma auditrix (Diptera: Sarcophagidae)

1. Acoustically guided movement in a three‐dimensional space is a complex behavioural task performed notably by birds, bats, and some insect species. The precision of acoustic orientation depends on the directionality of the hearing system as well as on auditory behaviour. 2. The fly Emblemasoma auditrix Diptera (Sarcophagidae) is a parasitoid of the cicada Okanagana rimosa Auchenorrhyncha (Cicadidae) and locates its host in the complex habitat of a forest. The phonotactic behaviour of the fly was analysed experimentally with emphasis on the vertical domain in the field. Different experimental setups allowed discriminating subsequent steps in the phonotactic behaviour of E. auditrix. 3. During the phonotactic flight, flies first landed on landmarks, which were used to re‐adjust to the elevation of the sound source. Acoustic targets were located from these resting positions. The sound source elevation was detected at the start of the flight as the longitudinal body axis was adjusted to the inclination of the target sound source. 4. Flies usually did not land directly upon the sound source, but landed nearby, and most often above the target. Within the target area, types of movement for the final approach differed in respect to target position; flies walked predominantly if the final target was located above or below, but for horizontally located targets much of the distance was covered by flight. 5. In conclusion, E. auditrix can locate the acoustic target in complex habitats and uses a flexible multi‐step approach for short‐range phonotaxis.     

The directionality of the ear of the pigeon (Columba livia)

Summary The directionality of cochlear microphonic potentials in the azimuthal plane was investigated in the pigeon (Columba livia), using acoustic free-field stimulation (pure tones of 0.25–6 kHz).At high frequencies in the pigeon's hearing range (4–6 kHz), changing azimuth resulted in a maximum change of the cochlear microphonic amplitude by about 20 dB (SPL). The directionality decreased clearly with decreasing frequency.Acoustic blocking of the contralateral ear canal could reduce the directional sensitivity of the ipsilateral ear by maximally 8 dB. This indicates a significant sound transmission through the bird's interaural pathways. However, the magnitude of these effects compared to those obtained by sound diffraction (maximum > 15 dB) suggests that pressure gradients at the tympanic membrane are only of subordinate importance for the generation of directional cues.The comparison of interaural intensity differences with previous behavioral results confirms the hypothesis that interaural intensity difference is the primary directional cue of azimuthal sound localization in the high-frequency range (2–6 kHz).Abbreviations CM cochlear microphonic potential - IID interaural intensity difference - IID-MRA minimum resolvable angle calculated from interaural intensity difference - MRA minimum resolvable angle - OTD interaural ongoing time difference - RMS root mean square - SPL sound pressure level     

Multiple paternity in the red-eyed treefrog Agalychnis callidryas (Cope)

DNA fingerprinting of tadpoles from two different ‘one female-two male’ matings of the red-eyed treefrog, Agalychnis callidryas, revealed multiple paternity of offspring. Offspring were assigned paternity based on bands shared with the putative father, but not shared between putative fathers or with the mother. Paternity was split 44/56% and 36/64% in two matings. These results do not support a hypothesis of sperm priority in access to unfertilized eggs by primary males. Multiple paternity may be commonplace in species of anurans with matings by multiple males.     

Multimodal signals in male European treefrog (Hyla arborea) and the influence of population isolation on signal expression

In nocturnal treefrogs, mate choice implies the use of acoustic and visual signals. Multimodality is suspected to have evolved for either information redundancy or information complementariness. It is essential to explore multimodality in a natural context to understand the selection pressures operating on the signals. In the present study, we investigated calling and coloration in relation to male biometry and condition in four populations of European treefrog (Hyla arborea) varying in size and genetic isolation. We compared the signal intensity between core and satellite populations to estimate the impact of genetic diversity on male secondary sexual traits. The results obtained show important regional variations in both traits, likely as a result of local adaptations. Call and coloration are weakly correlated within an individual, implying that these traits likely convey different information about the signaller's identity or quality, thus supporting the hypothesis of complementariness of multiple messages. By contrast to the experimental evidence, we find that call and coloration are not related to male condition (as estimated by the residual of mass over size), suggesting that the condition‐dependence of these traits may be mediated by complex mechanisms not accurately reflected by the chosen estimator. Finally, male call and colour phenotypes present no robust pattern of variation with isolation status, probably because of variation in local selective pressures and in history of population dynamics. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 633–647.     

Structure of the stridulatory apparatus and analysis of the sound produced by Smicronyx fulvus and Smicronyx sordidus (coleoptera,curculionidae, erirrhininae,smicronychini)

Wing folding spicules, elytral binding patches, and elytral locking devices of adult male and female seed weevils, Smicronyx fulvus LeConte and S. sordidus LeConte, involved in stridulation are described. Sound is produced by both sexes of the two species when the plectrum, paired conical teeth located along the anterior margin of the dorsally elevated seventh sternite, is struck against an elongate file, the pars stridens, on the under surface of the apical portion of each elytron. A second plectrum, on the sixth tergite, is well-developed in males of both species and is used by males to produce sound before and during mating. Sex-specific and species-specific differences in the sound produced is attributed to structural variation in the pars stridens and the elytra. The pars stridens determines frequencies, while the elytra may further modify the sound. The frequency range for male S. fulvus is 1,000 cycles per second (cps) through 13,000 cps and for male S. sordidus is 2,500 cps through 13,000 cps. The frequency range for female S. fulvus is 2,000 cps through 11,500 cps and for female S. sordidus is 900 cps through 11,500 cps.     

Localization dominance and the effect of frequency in the Mongolian Gerbil, Meriones unguiculatus

Due to its good low-frequency hearing, the Mongolian Gerbil (Meriones unguiculatus) has become a well-established animal model for human hearing. In humans, sound localization in reverberant environments is facilitated by the precedence effect, i.e., the perceptual suppression of spatial information carried by echoes. The current study addresses the question whether gerbils are a valid animal model for such complex spatial processing. Specifically, we quantify localization dominance, i.e., the fact that in the context of precedence, only the directional information of the sound which reaches the ear first dominates the perceived position of a sound source whereas directional information of the delayed echoes is suppressed. As localization dominance is known to be stimulus-dependent, we quantified the extent to which the spectral content of transient sounds affects localization dominance in the gerbil. The results reveal that gerbils show stable localization dominance across echo delays, well comparable to humans. Moreover, localization dominance systematically decreased with increasing center frequency, which has not been demonstrated in an animal before. These findings are consistent with an important contribution of peripheral-auditory processing to perceptual localization dominance. The data show that the gerbil is an excellent model to study the neural basis of complex spatial-auditory processing.     

Regeneration of axotomized tympanal nerve fibres in the adult grasshopper Chorthippus biguttulus (L.) (Orthoptera: Acrididae) correlates with regaining the localization ability

Adult males of the grasshopper Chorthippus biguttulus exhibit a stereotyped turning behaviour towards the direction of the female song. This behaviour has been used to study the regeneration of synaptic connections used for pattern recognition and sound localization. Unilaterally deafened animals are not able to localize the sound direction and turn exclusively towards the intact side, regardless of the speaker position. This behaviour does not change with postoperative time. After an axotomy of the tympanic nerve fibres the sensory axons regrow and regenerate their synaptic contacts which is deduced from the recovery of the ability to localize sound. The behavioural threshold for stimulation from the operated side is increased by approx. 5 dB SPL. The probability of correct turning towards the operated side increases with postoperative time. The ability for lateralization improves with postoperative time and may reach values of intact animals (discrimination of 1–2 dB SPL difference. Animals with two operations (axotomy of one tympanal nerve and blocking of the other ear) do not react to the female song, which suggests that recognition of the species-specific song pattern is not possible with the regenerated fibres and their synaptic connections alone. Neuroanatomical studies show that the regeneration of localization ability is correlated with an ingrowth of sensory fibres into the frontal auditory neuropil of the metathoracic ganglion.     

Species of the genus Psoroptes (Acari: Psoroptidae): A taxonomic consideration

The biosystematic status of mite species belonging to the genus Psoroptes Gervais, 1841 is difficult to determine by phenotypic methods and has been subject to taxonomic revisions and ongoing debate. At present, the existence of five species, P. cuniculi (Delafond, 1859), P. ovis (Hering, 1838), P. equi (Hering, 1838), P. cervinus Ward, 1915 and P. natalensis Hirst, 1919, is generally accepted. This classification is based mainly on the host species, the localization of the mites on their hosts and morphological characters of male mites. However, a critical review of the literature indicates that the features used to discriminate between the five species are not unequivocal: (a) the localization of mite populations on host animals is not completely strict, (b) the lengths of the outer opisthosomal setae of male mites, which are the main morphological features used for species discrimination, overlap between the five postulated species, and (c) host specificity cannot be deduced from results of transfer experiments. Rather, conspecificity of the members of the genus Psoroptes has to be presumed which is supported by molecular genetic analyses. On these grounds and on rules of priority P. cervinus Ward, 1915, P. cuniculi (Delafond, 1859), P. natalensis Hirst, 1919 and P. ovis (Hering, 1838) are seen as synonyms of P. equi (Hering, 1838).     

Azimuthal sound localization in the European starling (Sturnus vulgaris)

Summary The physical measurements reported here test whether the European starling (Sturnus vulgaris) evaluates the azimuth direction of a sound source with a peripheral auditory system composed of two acoustically coupled pressure-difference receivers (1) or of two decoupled pressure receivers (2).A directional pattern of sound intensity in the freefield was measured at the entrance of the auditory meatus using a probe microphone, and at the tympanum using laser vibrometry. The maximum differences in the soundpressure level measured with the microphone between various speaker positions and the frontal speaker position were 2.4 dB at 1 and 2 kHz, 7.3 dB at 4 kHz, 9.2 dB at 6 kHz, and 10.9 dB at 8 kHz. The directional amplitude pattern measured by laser vibrometry did not differ from that measured with the microphone. Neither did the directional pattern of travel times to the ear. Measurements of the amplitude and phase transfer function of the starling's interaural pathway using a closed sound system were in accord with the results of the free-field measurements.In conclusion, although some sound transmission via the interaural canal occurred, the present experiments support the hypothesis 2 above that the starling's peripheral auditory system is best described as consisting of two functionally decoupled pressure receivers.Abbreviations CM cochlear microphonics - ITD interaural time difference - IID interaural intensity difference - MRA minimum resolvable angle - dB SPL sound-pressure level (re 0.00002 Pa)     

Improvements of Sound Localization Abilities by the Facial Ruff of the Barn Owl (Tyto alba) as Demonstrated by Virtual Ruff Removal

Background

When sound arrives at the eardrum it has already been filtered by the body, head, and outer ear. This process is mathematically described by the head-related transfer functions (HRTFs), which are characteristic for the spatial position of a sound source and for the individual ear. HRTFs in the barn owl (Tyto alba) are also shaped by the facial ruff, a specialization that alters interaural time differences (ITD), interaural intensity differences (ILD), and the frequency spectrum of the incoming sound to improve sound localization. Here we created novel stimuli to simulate the removal of the barn owl''s ruff in a virtual acoustic environment, thus creating a situation similar to passive listening in other animals, and used these stimuli in behavioral tests.

Methodology/Principal Findings

HRTFs were recorded from an owl before and after removal of the ruff feathers. Normal and ruff-removed conditions were created by filtering broadband noise with the HRTFs. Under normal virtual conditions, no differences in azimuthal head-turning behavior between individualized and non-individualized HRTFs were observed. The owls were able to respond differently to stimuli from the back than to stimuli from the front having the same ITD. By contrast, such a discrimination was not possible after the virtual removal of the ruff. Elevational head-turn angles were (slightly) smaller with non-individualized than with individualized HRTFs. The removal of the ruff resulted in a large decrease in elevational head-turning amplitudes.

Conclusions/Significance

The facial ruff a) improves azimuthal sound localization by increasing the ITD range and b) improves elevational sound localization in the frontal field by introducing a shift of iso–ILD lines out of the midsagittal plane, which causes ILDs to increase with increasing stimulus elevation. The changes at the behavioral level could be related to the changes in the binaural physical parameters that occurred after the virtual removal of the ruff. These data provide new insights into the function of external hearing structures and open up the possibility to apply the results on autonomous agents, creation of virtual auditory environments for humans, or in hearing aids.     

The role of the medial septum in the acoustic trachea of the cricket Gryllus bimaculatus

The auditory organs of the cricket which are situated in the front legs are joined together by a large transverse trachea which decisively influences their directional characteristics. The transverse trachea is medially divided by a septum. The importance of this septum for the localization of a sound source was tested by means of behavioural experiments in which the phonotactic movements of intact Gryllus bimaculatus females were compared quantitatively with those of the same specimen after perforation of the septum. The septal perforation does not noticeably influence locomotion in the absence of acoustic stimuli but selectively changes essential characteristics of phono taxis: 1) The animals walk in less straight lines. The oscillations around the mean course, typical of phonotaxis, are increased in amplitude, while the frequency decreases. 2) Course deviations from the direction of the sound source become more pronounced. 3) The threshold for phonotaxis is raised by about 10 dB. 4) Both the speed at which the animals walk and the proportion of time during which they are mobile are reduced. The results are discussed in relation to the role of the septum in the mechanism of sound localization, and with regard to its possible importance for the recognition of acoustic patterns.     

Decisions during Phonotaxis in the Bushcricket Requena verticalis (Orthoptera: Tettigoniidae): Do Females Change Direction to Alternative Male Calls?

Female bushcrickets (Requena verticalis, Listroscelidinae, Tettigoniidae) show a preference between male calls that differ in three parameters, temporal structure, frequency and intensity. In a two-choice speaker situation they prefer louder calls, songs in which the upper part of the frequency spectrum occupies higher frequencies and calls with short chirps rather than those with longer chirps. In an experiment females were offered an alternative call while orienting to a model of their conspecific song. The alternative call was demonstrably preferred by females when presented in a paired-speaker trial. Two motivational states were identified where females moved slowly or quickly to a sound source. Movement patterns were associated with the form of stimulus and slow females were discriminatory of male calls while fast females were not. Slow females changed speaker preference when the alternate speaker broadcast a call model with the higher frequency peak or the song model with the shorter chirps. By comparison fast moving females only changed course when the alternative sound source was louder.     

Precognitive and cognitive elements in sound localization

Sound localization behavior is of great importance for an animal's survival. To localize a sound, animals have to detect a sound source and assign a location to it. In this review we discuss recent results on the underlying mechanisms and on modulatory influences in the barn owl, an auditory specialist with very well developed capabilities to localize sound. Information processing in the barn owl auditory pathway underlying the computations of detection and localization is well understood. This analysis of the sensory information primarily determines the following orienting behavior towards the sound source. However, orienting behavior may be modulated by cognitive (top-down) influences such as attention. We show how advanced stimulation techniques can be used to determine the importance of different cues for sound localization in quasi-realistic stimulation situations, how attentional influences can improve the response to behaviorally relevant stimuli, and how attention can modulate related neural responses. Taken together, these data indicate how sound localization might function in the usually complex natural environment.     

Properties of low-frequency head-related transfer functions in the barn owl (Tyto alba)

The barn owl (Tyto alba) possesses several specializations regarding auditory processing. The most conspicuous features are the directionally sensitive facial ruff and the asymmetrically arranged ears. The frequency-specific influence of these features on sound has consequences for sound localization that might differ between low and high frequencies. Whereas the high-frequency range (>3 kHz) is well investigated, less is known about the characteristics of head-related transfer functions for frequencies below 3 kHz. In the present study, we compared 1/3 octaveband-filtered transfer functions of barn owls with center frequencies ranging from 0.5 to 9 kHz. The range of interaural time differences was 600 μs at frequencies above 4 kHz, decreased to 505 μs at 3 kHz and increased again to about 615 μs at lower frequencies. The ranges for very low (0.5–1 kHz) and high frequencies (5–9 kHz) were not statistically different. Interaural level differences and monaural gains increased monotonically with increasing frequency. No systematic influence of the body temperature on the measured localization cues was observed. These data have implications for the mechanism underlying sound localization and we suggest that the barn owl’s ears work as pressure receivers both in the high- and low-frequency ranges.     

The function of sound in male spacing behaviour in bush-crickets (Tettigoniidae,Orthoptera)

This paper examines the spatial dynamics of an aggregation of singing male bush-crickets (Tettigoniidae) in respect to the degree of regularity between males. Our results establish that singing males of the genus Mygalopsis are regularly spaced within aggregations and that sound is involved in spacing. Attention is focussed on the male strategy of regular spacing within a widely dispersed aggregation in order to attract females reaching sexual maturity within the confines of that population. This is seen as an alternate to male aggregations attracting females from outside their boundary.     

Phonotaxis in the painted reed frog (Hyperolius marmoratus)

Summary A detailed study was conducted of the three-dimensional accuracy of phonotaxis by femaleHyperolius marmoratus. This analysis involved videotape recordings of phonotactic approaches to an elevated loudspeaker through a three-dimensional grid. Females readily resolved the sound source elevation, but the jump error angles describing the precision of approach were considerably greater in this three-dimensional analysis than in the more conventional two-dimensional ground approach analysis. Extensive use was made of visual cues in elevated phonotactic approach and lateral head scanning prior to jumps, often accompanied by vertical changes in head orientation, was frequent. The ability of such small anurans to localize a sound source in both the horizontal and vertical plane is remarkable.On leave from the Section of Neurobiology and Behavior, Cornell University, Ithaca, New York 14853, USA     

Ultrasonic courtship song of the yellow peach moth, <Emphasis Type="Italic">Conogethes punctiferalis</Emphasis> (Lepidoptera: Crambidae)

Although generation of ultrasound during courtship has been reported for an increasing number of moth species, the effect of the ultrasound on mating remains uncertain in many cases because of a lack of proper verification. Here we report that males of the yellow peach moth Conogethes punctiferalis (Crambidae) sexually communicate with females by emitting loud ultrasound (103 dB peak equivalent sound pressure level at 1 cm; dominant frequency 82 kHz) before attempting copulation. The male ultrasound consists of consecutive clicks (pulses) in the early phase of the sound train and consecutive pulses (burst) in the late phase. When females were deafened by puncturing the abdominal tympanic membranes, copulation never occurred. We found that deafened females did not assume the wing-raising posture, which, for normal pairs, always precedes successful copulation. Our findings indicate that male courtship ultrasound evokes wing-raising as an acceptance behavior from females, which in turn evokes a copulation attempt by a male.     

Effects of seawater temperature on sound characteristics in Ophidion rochei (Ophidiidae)

Although the sound production mechanisms of male and female Ophidion rochei (Ophidiidae) differ significantly, temperature affects them in the same manner. In both sexes, temperature correlated negatively with pulse period and positively with sound frequencies but had no, or weak effects on other sound characteristics.