Broadband Calls Research Articles

Representation of species-specific vocalizations in the medial geniculate body of the guinea pig

Individual nuclei of the auditory pathway contribute in a specific way to the processing of complex acoustical signals. We investigated the responses of single neurons to typical guinea pig vocalizations (purr, chutter, chirp and whistle) in the ventral part of the medial geniculate body (MGB) of anesthetized guinea pigs. The neuronal and population peristimulus time histograms (PSTHs) reflected the repetition frequency of individual phrases in the calls. The patterns of PSTHs correlated well with the sound temporal envelope in calls with short phrases (purr, chirp). The dominant onset character of the neuronal responses resulted in a lower correlation between the sound envelope and the PSTH pattern in the case of longer calls (chutter and whistle). A time-reversed version of whistle elicited on average a 13% weaker response than did the natural whistle. The rate-characteristic frequency (CF) profile provided only a coarse representation of the sound frequency spectrum without detailed information about the individual spectral peaks and their relative magnitudes. In comparison with the inferior colliculus (Suta et al. in J Neurophysiol 90:3794-3808, 2003), the processing of species-specific vocalizations in the MGB differs in: (1) a less precise representation of the temporal envelope in the case of longer calls, but not in the case of calls consisting of one or more short phrases; (2) a less precise rate-CF representation of the spectral envelope in the case of low-frequency calls, but not in the case of broad-band calls; (3) a smaller difference between the responses to natural and time-reversed whistle.

Psychoacoustic sampling as a reliable, non-invasive method to monitor orthopteran species diversity in tropical forests

We evaluated trained listener—based acoustic sampling as a reliable and non-invasive method for rapid assessment of ensiferan species diversity in tropical evergreen forests. This was done by evaluating the reliability of identification of species and numbers of calling individuals using psychoacoustic experiments in the laboratory and by comparing psychoacoustic sampling in the field with ambient noise recordings made at the same time. The reliability of correct species identification by the trained listener was 100 % for 16 out of 20 species tested in the laboratory. The reliability of identifying the numbers of individuals correctly was 100% for 13 out of 20 species. The human listener performed slightly better than the instrument in detecting low frequency and broadband calls in the field, whereas the recorder detected high frequency calls with greater probability. To address the problem of pseudoreplication during spot sampling in the field, we monitored the movement of calling individuals using focal animal sampling. The average distance moved by calling individuals for 17 out of 20 species was less than 1.5 m in half an hour. We suggest that trained listener—based sampling is preferable for crickets and low frequency katydids, whereas broadband recorders are preferable for katydid species with high frequency calls for accurate estimation of ensiferan species richness and relative abundance in an area.

The influence of wing morphology and echolocation on the gleaning ability of the insectivorous bat Myotis tricolor

On the basis of its external morphology, Myotis tricolor (Temminck, 1832) should be able to both aerial-feed and glean. Furthermore, this bat is known to use broadband calls of short duration, reinforcing the prediction that it gleans. However, results from this study indicate that M. tricolor does not commonly glean. This conclusion was reached after studying the foraging behaviour of M. tricolor in a flight room. We presented M. tricolor with mealworms, moths, mole crickets, beetles, and cicadas in a variety of ways that required either gleaning and (or) aerial feeding. Although M. tricolor readily took tethered prey, it did not take any of the variety of insects presented to it in a manner that required gleaning. We therefore compared its wing morphology and echolocation calls with those of several known gleaners, Nycteris thebaica E. Geoffroy, 1818, Myotis lucifugus (Le Conte, 1831), and Myotis septentrionalis (Trouessart, 1897), and an aerial forager, Neoromicia capensis (A. Smith, 1829). In a discriminant analysis wing-tip shape was the only variable to provide some degree of discrimination between species, with M. tricolor having more pointed wing tips than the known gleaners. Discriminant analysis of echolocation-call parameters grouped M. tricolor with the other Myotis species and separated it from N. capensis and N. thebaica. However, M. tricolor did not use harmonics as did the other Myotis species. The apparent failure of M. tricolor to glean might therefore be due to its relatively pointed wings and narrow-bandwidth echolocation calls, owing to the absence of harmonics in its calls.

Localization of aerial broadband noise by pinnipeds.

Although many pinnipeds (seals, sea lions, and walruses) emit broadband calls on land as part of their communication system, few studies have addressed these animals' ability to localize aerial broadband sounds. In this study, the aerial sound localization acuities of a female northern elephant seal (Mirounga angustirostris), a male harbor seal (Phoca vitulina), and a female California sea lion (Zalophus californianus) were measured in the horizontal plane. The stimulus was broadband white noise that was band pass filtered between 1.2 and 15 kHz. Testing was conducted in a hemi-anechoic chamber using a left/right forced choice procedure to measure the minimum audible angle (MAA) for each subject. MAAs were defined as half the angular separation of two sound sources bisected by a subject's midline that corresponded to 75% correct discrimination. MAAs were 4.7 degrees, 3.6 degrees, and 4.2 degrees for the northern elephant seal, harbor seal, and California sea lion, respectively. These results demonstrate that individuals of these pinniped species have sound localization abilities comparable to the domestic cat and rhesus macaque. The acuity differences between our subjects were small and not predicted by head size. These results likely reflect the relatively acute general abilities of pinnipeds to localize aerial broadband signals.

Function of male song in the greater white-lined bat, Saccopteryx bilineata

The use of vocalizations by males to attract mates and defend territories is widespread in birds but uncommon in mammals. In the greater white-lined bat, males defend small territories in tree buttresses and produce complex audible vocalizations. During the day, females roost within these territories, and males with females in their territories have higher reproductive success than males without females. To determine call function, we recorded male vocalizations on the island of Trinidad and made behavioural observations of the focal male and nearby bats at the time the calls were given. Noisy broadband calls were directed mainly at other males whereas tonal calls were used primarily in interactions with females. Two other types of calls were given independently of any observable context and may be involved in territorial defence or conspecific notification. Poisson regression showed that males with song repertoires that contained more unique types of composite syllables had more females in their territories. In addition, several acoustic features of one common call type significantly correlated with the number of females on a territory. Male vocalizations may, therefore, transfer information to colony members about male quality or fighting ability.

Sound localization of aerial broadband noise in pinnipeds

Pinnipeds (seals, sea lions, and walruses) emit broadband calls on land as part of their communication system in order to coordinate their reproductive activities. How well do they localize these types of signals? In this study, the aerial sound localization acuities of a harbor seal (Phoca vitulina), a California sea lion (Zalophus californianus), and a northern elephant seal (Mirounga angustirostris) were measured in the horizontal plane with a broadband white noise stimulus. Testing was conducted in a hemi-anechoic chamber using a left/right forced choice procedure to measure the minimum audible angle (MAA) for each subject. MAAs were defined as half the angular separation of two sound sources relative to a subject’s midline that corresponded to 75% correct discrimination. MAAs were 3.6, 4.2, and 4.7 deg for the harbor seal, California sea lion, and northern elephant seal, respectively. These results demonstrate that these pinniped species had sound localization abilities comparable to the domestic cat and rhesus macaques. The acuity differences between our subjects were small, were not predicted by head size, and therefore likely reflect the relatively acute abilities of other pinniped species to localize aerial broadband signals.

Laryngeal biomechanics and vocal communication in the squirrel monkey (Saimiri boliviensis).

The larynges of eight squirrel monkeys were harvested, dissected, mounted on a pseudotracheal tube, and phonated using compressed air. Patterns of vocal fold oscillation were compared with sound spectrograms of calls recorded from monkeys in our colony. Four different regimes of vocal fold activation were identified. Regime 1 resembled typical human vowel production, with regular vocal-fold vibration, a prominent fundamental frequency, and an accompanying series of harmonic overtones. This regime is likely to give rise to squirrel monkey "cackles," as well as a variety of other harmonically structured calls. In regime 2, the pattern of vibrations exhibited the presence of two or more unrelated frequencies (biphonation). This regime of glottal activity resembled the biphonation observed in many exemplars of "twitter" and "kecker" calls. The vocal folds oscillated continuously in regime 3, but produced glottal pulses whose amplitudes waxed and waned rhythmically. This phenomenon resulted in the percept of a series of discrete pulses, and may give rise to "errs," "churrs," and other calls composed of a rapid sequence of acoustic elements. In regime 4, the period of each oscillation was quasi-irregular. Shrieks and other broadband calls or call elements that lack an apparent fundamental frequency may be produced in this manner.

Evidence for an analytic perception of multiharmonic sounds in the bat, Megaderma lyra, and its possible role for echo spectral analysis.

For echolocation, the gleaning bat Megaderma lyra relies on short and broadband calls consisting of multiple harmonic components, each of which is downward frequency modulated. The harmonic components in M. lyra's calls have a relatively small frequency excursion and do not overlap spectrally. Broadband calls of other bat species, on the other hand, often consist of only a few harmonics which are modulated over broad and sometimes overlapping frequency ranges. A call consisting of narrow and nonoverlapping harmonic components may provide a less complete representation of target structure than a call which consists of broadly modulated components. However, a multiharmonic call may help the bats to perceive local spectral changes in the echo from shifts in the peak frequencies of single harmonics, and thereby to extract additional information about the target. To assess this hypothesis, the accuracy with which M. lyra can analyze frequency shifts of single partials in multiharmonic complex tones was investigated. A two-alternative, forced-choice behavioral task was used to measure M. lyra's frequency discrimination threshold for the third partial in complex tones whose spectral composition resembled that of the bat's sonar calls. The discrimination threshold for the third partial in a 21.5-kHz harmonic tone amounted to about 2% and was similar to the bat's pure-tone discrimination threshold at 64.5 kHz. Discrimination performance was essentially unaffected by random frequency changes of the other partials and by reducing stimulus duration from 50.5 to 1.5 ms. Both findings are in accordance with predictions made on the basis of the shape of M. Ivra's cochlear filters. The comparison between the observed frequency discrimination performance and a computational estimate of the expected frequency shift in the third harmonic of an echo reflected by a simple, two-front target showed that M. lyra's frequency resolution is sufficient for analyzing the target-specific information conveyed by shifts in the peak frequency of single echo components.

A distributed dynamic channel assignment method for a mixture of narrowband and broadband traffic in a cellular system

This paper proposes an autonomous distributed dynamic channel assignment method, considering a mixture of narrowband and broadband traffic processing in a multicarrier TDMA mobile communication system. As the first step, narrowband and broadband calls are considered, and the system in which the calls share the carrier [carrier-sharing (CS) method], and the system in which the calls use separate carriers [carrier partitioning (CP) method], are compared. For each method, a mathematical model is proposed for the blocking probability analysis in the dynamic channel assignment, based on the steady-state probability. Using the models, the range for which the CS method exhibits a better performance than the CP method is specified. A problem in the dynamic channel assignment in the CS method is that the transmission quality of the broadband call may be degraded by interference from a narrowband call. In order to overcome such a problem, a method is proposed where the suppression of the leftovers of nonavailable channels for assignment, the multicarrier modulation using frequency hopping, the reallocation connection of calls, the variable-rate transmission connection, and the retry-type hand-off are combined. It is verified by computer simulation that the proposed method exhibits a performance which is better than that of the conventional method. © 2000 Scripta Technica, Electron Comm Jpn Pt 1, 83(11): 85–94, 2000

Moth hearing in response to bat echolocation calls manipulated independently in time and frequency.

We measured the auditory responses of the noctuid moth Noctua pronuba to bat echolocation calls which were manipulated independently in time and frequency. Such manipulations are important in understanding how insect hearing influences the evolution of echolocation call characteristics. We manipulated the calls of three bat species (Rhinolophus hipposideros, Myotis nattereri and Pipistrellus pipistrellus) that use different echolocation call features by doubling their duration or reducing their frequency, and measured the auditory thresholds from the A1 cells of the moths. Knowing the auditory responses of the moth we tested three predictions. (i) The ranking of the audibility of unmanipulated calls to the moths should be predictable from their temporal and/or frequency structure. This was supported. (ii) Doubling the duration of the calls should increase their audibility by ca. 3 dB for all species. Their audibility did indeed increase by 2.1-3.5 dB. (iii) Reducing the frequency of the calls would increase their audibility for all species. Reducing the frequency had small effects for the two bat species which used short duration (2.7-3.6 ms) calls. However, the relatively long-duration (50 ms), largely constant-frequency calls of R. hipposideros increased in audibility by 21.6 dB when their frequency was halved. Time and frequency changes influence the audibility of calls to tympanate moths in different ways according to call design. Large changes in frequency and time had relatively small changes on the audibility of calls for short, largely broadband calls. Channelling energy into the second harmonic of the call substantially decreased the audibility of calls for bats which use long-duration, constant-frequency components in echolocation calls. We discuss our findings in the contexts of the evolution of both bat echolocation call design and the potential responses of insects which hear ultrasound.

One tone, two ears, three dimensions: a robotic investigation of pinnae movements used by rhinolophid and hipposiderid bats.

Bats, which echolocate using broadband calls, are believed to employ the passive acoustic filtering properties of the head and pinnae to provide spectral cues which encode 3-D target angle. Microchiropteran species whose calls consist of a single, constant frequency harmonic (i.e., some species in the families Rhinolophidae and Hipposideridae) may create additional acoustic localization cues via vigorous pinna movements. In this work, two types of echolocation cues generated by moving a pair of receivers aboard a model sensor head are investigated. In the first case, it is supposed that a common 3-D echolocation principle employed by all bats is the creation of alternative viewing perspectives, and that constant frequency (CF) echolocators use pinna movement rather than morphology to alter the acoustic axes of their perceptual systems. Alternatively, it is possible rhinolophids and hipposiderids move their ears to create dynamic cues--in the form of frequency and amplitude modulations--which vary systematically with target elevation. Here the use of binaural and monaural timing cues derived from amplitude modulated echo envelopes are investigated. In this case, pinna mobility provides an echolocator with a mechanism for creating dramatic temporal cues for directional sensing which, unlike interaural timing differences, do not degrade with head size.

Greater spear-nosed bats discriminate group mates by vocalizations

Individuals often benefit from identifying their prospective social partners. Some species that live in stable social groups discriminate between their group mates and others, basing this distinction on calls that differ among individuals. Vocalizations that differ between social groups are much less common, and few studies have demonstrated that animals use group-distinctive calls to identify group mates. Female greater spear-nosed bats,Phyllostomus hastatus, live in stable groups of unrelated bats and give audible frequency, broadband calls termed screech calls when departing from the roost and at foraging sites. Previous field observations suggested that bats give screech calls to coordinate movements among group members. Prior acoustic analyses of 12 acoustic variables found group differences but not individual differences. Here, we use the same acoustic variables to compare calls from three cave colonies, and find that calls differ between caves. We also report results from field and laboratory playback experiments designed to test whether bats use acoustic differences to discriminate calls from different colonies, groups or individuals. Results from field playbacks indicate that response depends on the cave of origin, indicating that bats can discriminate among calls from different caves. This discrimination ability may be based, in part, on whether calls are familiar or unfamiliar to the listening bats. Laboratory playbacks demonstrate that bats discriminate calls given by their group mates from calls given by other bats from the same cave irrespective of familiarity. However, these experiments provide no evidence that bats discriminate among individuals. Previous field work indicates that females that forage with social group mates may benefit from shared information about food or mutual defence of feeding sites. Indicating group membership is essential, since these benefits appear to be restricted to group mates.

Vocal learning by greater spear-nosed bats.

Vocal learning is well known among passerine and psittacine birds, but most data on mammals are equivocal. Specific benefits of vocal learning are poorly understood for most species. One case where vocal learning should be favoured by selection is where calls indicate group membership and group mates are unrelated. Female greater spear-nosed bats, Phyllostomus hastatus, live in stable groups of unrelated bats and use loud, broadband calls to coordinate foraging movements of social group mates. Bats benefit from group foraging. Calls differ between female social groups and cave colonies, and playback experiments demonstrate that bats perceive these acoustic differences. Here I show that the group distinctive structure of calls arises through vocal learning. Females change call structure when group composition changes, resulting in increased similarity among new social group mates. Comparisons of transfers with age-matched half-sibs indicate that call changes are not simply due to maturation, the physical environment or heredity. These results suggest that studies testing vocal learning in mammals could profit by focusing on vocalizations that signify group membership.

Acoustic discrimination of passerine anti-predator signals by Australian raptors

Acoustic discrimination of anti-predator calls was examined in 11 species of Australian raptors, including 5 Falco species and 2 species of Elanus kites, by their responses to the playback of alarm and distress calls of Australian passerines. The present study investigated the ability of raptors to discriminate between alarm and distress calls that have different acoustic properties and are emitted in different behavioural contexts. The raptors were tested with broad-band calls (containing a wide range of frequencies) given as distress calls, mobbing calls and alarm calls to terrestrial predators, and with narrow-band calls (comprising a narrow range of frequencies) typically given as a response to flying predators. Raptor responses were categorised into three classes based on head orientation (or lack thereof) towards the sound source (i.e. one of 2 or 4 speakers positioned in the cage set-up); (1) ‘correct response’ – the raptor looked directly at the speaker; (2) ‘incorrect response’ – the raptor detected the sound, but oriented the head in a direction other than towards the sound source; (3) ‘no response’. All raptor species showed a higher percentage of correct responses (60–100%) for broad-frequency vocalisations and a lower percentage of correct responses (usually 0–40%) and more incorrect responses for narrow-band vocalisations. Further, all raptors showed a greater rate of overall responsiveness to broad-band alarm and distress calls than narrow-band calls, indicating a higher interest level in the former. The behavioural implications of acoustic discrimination by Australian raptors to different types of alarm call are discussed.

Target localization aboard a tone emitting robotic echolocation system with mobile receivers

Using only two receivers, animals are able to determine the location of a target in three dimensions. Echolocating bats, which employ broadband calls, have access to spectral cues that may be used, in conjunction with interaural disparities of intensity (IIDs) and/or time (ITDs), to encode a 3-D target angle. This work addresses the question: what cues might a tone-emitting echolocator employ to determine target azimuth and elevation? To this end, a 6-degree-of-freedom robotic sensor head was built to investigate the direction cues that might be generated by the systematic pinnae scanning movements used by many species of tone emitting bats. The first strategy investigated was the use of receiver motion to rotate the SONAR horizon through discrete IID sampling positions so that a 3-D target angle can be resolved across IID readings taken at two or more receiver positions. Next, the change in amplitude measured by continuously scanning pinnae was used to create temporal cues, which vary systematically with the target angle. In this scheme, angular resolution depends upon scan speed (ms) rather than interaural separation (microsecond). Finally, the use of Doppler shifts to encode the target angle through the cosine law was investigated.

Morphology, Karyology, and Echolocation Calls of Rhogeessa (Chiroptera: Vespertilionidae) from the Yucatan Peninsula

Morphological analyses of 23 specimens of Rhogeessa from localities in the Yucatan Peninsula revealed significant sexual dimorphism in some characters. Although there was no significant sexual dimorphism in length of ear or length of tragus, males had conspicuous ear glands and females did not. Rhogeessa produced short, broad-band echolocation calls that showed significant inter-individual variation. Karyotypes of 16 Rhogeessa were similar to a karyotype reported from Belize and different than those of R. tumida from areas adjacent to the Yucatan Peninsula. Yucatan populations also are morphologically distinctive and we assign them to a separate species, R. aeneus Goodwin.

Regulation of rat maternal behavior by broadband pup vocalizations

Pups emit a broadband vocalization ranging from 2 to 40 kHz that generally occurs when the mother sits or steps on her pups. In an earlier study (Ihnat, White, and Barfield, under review), we noted that the mother was more likely to move on the pups and step on them when she was temporarily deafened with ear plugs. This research was extended in the present study by using two different methods to prevent 2-day-old pups from emitting broadband vocalizations. In the first experiment, the female was presented with six of her pups for a 10-min test. In half the tests, the pups had been anesthetized with pentobarbital; in the other half, the pups received saline. In the second experiment, females were presented with pups whose mouths had been sealed with an adhesive substance. In a final study, pups received the adhesive substance on their ventral surface as a control. When the pups were unable to vocalize, there was an increase in the amount of time per bout that the mother spent in incidental contact with her pups; in the sham condition, however, there was no change in her behavior. In addition, the female was less likely to group all her pups within the 10-min duration of the test when the pups could not call. Since pups emit primarily broadband vocalizations in response to handling prior to 4 days of age, broadband calls appear to reduce incidental contact and facilitate retrieval into a group.

The echolocation calls of hoary (Lasiurus cinereus) and silver-haired (Lasionycteris noctivagans) bats as adaptations for long- versus short-range foraging strategies and the consequences for prey selection

Amongst aerial-feeding insectivorous bats, differences in the design of echolocation calls appear to be associated with differences in foraging strategy. Recordings and observations of hoary (Lasiurus cinereus) and silver-haired (Lasionycteris noctivagans) bats in Manitoba, Canada, support such an association. Lasionycteris noctivagans use multiharmonic search–approach calls with an initial frequency sweep and a constant frequency tail. Such calls are suited for bats foraging in the open but near obstacles, and pursuing prey detected at relatively close range. This is the foraging strategy employed by this relatively slow, manoeuverable species. Lasiurus cinereus employ single harmonic search–approach calls that are low (20–17 kHz), essentially constant frequency signals. Calls of this design are suited for long-range target detection in open air situations, the foraging strategy used by L. cinereus. Differences in call design may explain dietary differences between the two species. Lasiurus cinereus consistently prey on large insects. The low, constant frequency design of their calls means that small insects are detectable only at close range and are thus difficult for this fast-dying bat to catch. The broad-band calls used by L. noctivagans do not restrict prey detection and these bats prey on a wider range of insects. Similar restrictions on prey detection, caused by echolocation call specializations, may be important in producing what might otherwise be considered active prey selection by some insectivorous bats.