The Energetics of Acoustic Signaling in Anurans and Insects

SYNOPSIS. This review focuses on the energetics of advertisementcalls in frogs and insects (mostly ensiferans). I also reviewa number of methodological questions relating to the most appropriateways to normalize metabolic rates for calling animals and tocalculate efficiency of sound production Although the mechanism of sound production is very differentin these groups (vocalization vs. stridulation), net metaboliccosts normalized to mass are similar among species producingthe most conspicuous calls. Features of the call that interactto determine energetic cost include repetition rate, call duration,and intensity (sound pressure level). Anurans tend to producelouder (more intense) calls while ensiferans tend to producesound during a greater proportion of the calling bout. All evidencesuggests that advertisement calls are produced aerobically andthe aerobic costs are similar or exceed the aerobic (but notnecessarily total costs) of terrestrial locomotion The pattern of radiated sound tends to be constant within aspecies and can be predicted to some degree from characteristicsof an animal's acoustic radiator. Efficiency of sound production(acoustic power/net metabolic power) is low (0.05 to 6%) andvariable when compared to locomotion ({small tilde} 10–20%).From the present sample it appears that frogs are more efficientthan ensiferans, but as more katydids are studied this trendmay not hold. Of the factors that have been identified as determiningefficiency the most important are the match between the sizeof the radiator and wavelength radiated, the absorption propertiesof the environment immediately around the animal, and the presenceor absence of structures such as baffles and acoustic burrows     

Call Transmission Efficiency in Native and Invasive Anurans: Competing Hypotheses of Divergence in Acoustic Signals

Invasive species are a leading cause of the current biodiversity decline, and hence examining the major traits favouring invasion is a key and long-standing goal of invasion biology. Despite the prominent role of the advertisement calls in sexual selection and reproduction, very little attention has been paid to the features of acoustic communication of invasive species in nonindigenous habitats and their potential impacts on native species. Here we compare for the first time the transmission efficiency of the advertisement calls of native and invasive species, searching for competitive advantages for acoustic communication and reproduction of introduced taxa, and providing insights into competing hypotheses in evolutionary divergence of acoustic signals: acoustic adaptation vs. morphological constraints. Using sound propagation experiments, we measured the attenuation rates of pure tones (0.2–5 kHz) and playback calls (Lithobates catesbeianus and Pelophylax perezi) across four distances (1, 2, 4, and 8 m) and over two substrates (water and soil) in seven Iberian localities. All factors considered (signal type, distance, substrate, and locality) affected transmission efficiency of acoustic signals, which was maximized with lower frequency sounds, shorter distances, and over water surface. Despite being broadcast in nonindigenous habitats, the advertisement calls of invasive L. catesbeianus were propagated more efficiently than those of the native species, in both aquatic and terrestrial substrates, and in most of the study sites. This implies absence of optimal relationship between native environments and propagation of acoustic signals in anurans, in contrast to what predicted by the acoustic adaptation hypothesis, and it might render these vertebrates particularly vulnerable to intrusion of invasive species producing low frequency signals, such as L. catesbeianus. Our findings suggest that mechanisms optimizing sound transmission in native habitat can play a less significant role than other selective forces or biological constraints in evolutionary design of anuran acoustic signals.     

Sensory Motor Interfacing in Acoustic Behavior of Anurans

SYNOPSIS. Anurans respond to different acoustic signals in adistinct manner. Moreover, acoustic behavior strongly dependson the inner state of the animal and the social context. Neuroanatomicalstudies as well as extra- and intracellular recordings werecarried out to examine the problem of audio-motor interfacing.Acoustic signals are processed in a partly hierarchical andpartly distributed manner in the anuran central nervous system.Features are represented in a topographical manner. Auditorymaps in the midbrain torus semicircularis are read by a subsetof neurons of the laminar and magnocellular subnuclei, whichinterconnect the auditory pathway and premotor centres. These,in turn, feed into neural networks controlling vocalization,locomotion, or autonomic functions. Our data on the cytoarchitectonicorganization and connectivity of the interfacing neurons giverise to a population coding hypothesis which may explain thedifferential evaluation of acoustic signals. Additionally, immunohistochemicalfindings reveal strong neuromodulatory and hormonal input, especiallyonto interfacing neurons, suggesting intense influence on thephysiological properties of these cells.     

Character Displacement and Acoustic Insects

Crickets, katydids, grasshoppers, and cicadas should be idealfor illustrating character displacement. Their species-specificcalling songs are of direct importance to reproductive success,and species with similar songs and overlapping ranges shoulddevelop greater differences in their songs in areas of sympatry.Although many pairs of species have suitably similar songs andoverlapping ranges and the data concerning songs are extensivefor crickets and katydids, no convincing example of characterdisplacement has been found in acoustic insects. Lack of suchexamples can be attributed to false premises, to divergencecompleted in allopatry. to elimination of geographical variationin song by gene flow between the zone of sympatry and the zonesof allopatry, and to the scanty sample that has been intensivelystudied.     

Synchronous and Alternating Choruses in Insects and Anurans: Common Mechanisms and Diverse Functions

SYNOPSIS. Neighboring males in rhythmically calling insectsand anurans often chorus in a synchronizing or alternating fashion.Neuroethological investigations of chorusing species revealthat their rhythms are maintained by pacemakers and that a basicinteractive algorithm, common to many species, yields the collectivesynchrony or alternation observed. Traditionally, synchronyhas been viewed as a cooperative event. However, recent evidencesuggests that a collective synchronous display can also be anincidental outcome of signal "jamming" activities between neighboringmales competing to attract females. This arises when femalephonotaxis is influenced by a precedence effect in which thefirst of two or more closely synchronized calls is preferred.Under such circumstances, males are selected to adopt a timingmechanism averting following calls. If males happen to callat comparable rates, the adopted mechanism can yield synchronyas a by-product. Alternation, too, may be produced by a similarmechanism and also represent an epiphenomenon. That alternation,as opposed to synchrony, results may be a mere artefact of thespecies' solo calling rate, but perceptual constraints may selectspecifically for alternation in some species.     

Selective and Efficient Neural Coding of Communication Signals Depends on Early Acoustic and Social Environment

Previous research has shown that postnatal exposure to simple, synthetic sounds can affect the sound representation in the auditory cortex as reflected by changes in the tonotopic map or other relatively simple tuning properties, such as AM tuning. However, their functional implications for neural processing in the generation of ethologically-based perception remain unexplored. Here we examined the effects of noise-rearing and social isolation on the neural processing of communication sounds such as species-specific song, in the primary auditory cortex analog of adult zebra finches. Our electrophysiological recordings reveal that neural tuning to simple frequency-based synthetic sounds is initially established in all the laminae independent of patterned acoustic experience; however, we provide the first evidence that early exposure to patterned sound statistics, such as those found in native sounds, is required for the subsequent emergence of neural selectivity for complex vocalizations and for shaping neural spiking precision in superficial and deep cortical laminae, and for creating efficient neural representations of song and a less redundant ensemble code in all the laminae. Our study also provides the first causal evidence for ‘sparse coding’, such that when the statistics of the stimuli were changed during rearing, as in noise-rearing, that the sparse or optimal representation for species-specific vocalizations disappeared. Taken together, these results imply that a layer-specific differential development of the auditory cortex requires patterned acoustic input, and a specialized and robust sensory representation of complex communication sounds in the auditory cortex requires a rich acoustic and social environment.     

Acoustic Analysis of Signals in Mammals

Spectral-prosodic characteristics of signals were analyzed synchronously with the main frequency for 32 mammals and one reptile using a KAZ-01 complex sound analyzer. The method of phoneme analysis was adapted for analysis of the signals. The main features of signaling in mammals have been revealed, which differed from those in birds and humans: low frequency range, low fundamental frequency, small number of local maxima, small frequency distance between them, and a significant amount of local maxima with zero energy.     

From Sender to Receiver: Propagation and Environmental Effects on Acoustic Signals

SYNOPSIS. Acoustic signals transmitted over large distancesdiffer significantly from those emitted by the signaler. Acousticsignals degrade in amplitude, spectral and temporal structureas they propagate through theenvironment. A great deal of workon acoustic communication is aimed at understanding the selectiveforces imposed by the environment on animal signals. I willdiscuss the physical constraints the environment puts on acousticcommunication, and then discuss similarities in communicationby anurans and insects that relate these environmental constraintsto their signaling systems. Lastly, I show how changes in signalsduring propagation relate to changes in signal perception duringphonotaxis, and thus, how propagation relates to mate choiceand sexual selection     

Acoustic Signals in Mouse Maternal Behavior: Retrieving and Cannibalism

The purpose of this study was to show that juvenile acoustic signals are not essential prerequisites for inhibiting parental aggression or for eliciting maternal care. This was demonstrated by comparing the behavior of Rb-3 and GFF+/+ female mice with normal audition and that of a deaf mutant GFF dn/dn. 243 ♀♀ and 609 litters were used to measure cannibalism and 54 ♀♀ to measure speed of retrieval. Neither rate of cannibalism nor speed of retrieval were modified by deafness, thus verifying the proposed hypothesis.     

The Acoustic Structure and Information Content of Female Koala Vocal Signals

Determining the information content of animal vocalisations can give valuable insights into the potential functions of vocal signals. The source-filter theory of vocal production allows researchers to examine the information content of mammal vocalisations by linking variation in acoustic features with variation in relevant physical characteristics of the caller. Here I used a source-filter theory approach to classify female koala vocalisations into different call-types, and determine which acoustic features have the potential to convey important information about the caller to other conspecifics. A two-step cluster analysis classified female calls into bellows, snarls and tonal rejection calls. Additional results revealed that female koala vocalisations differed in their potential to provide information about a given caller’s phenotype that may be of importance to receivers. Female snarls did not contain reliable acoustic cues to the caller’s identity and age. In contrast, female bellows and tonal rejection calls were individually distinctive, and the tonal rejection calls of older female koalas had consistently lower mean, minimum and maximum fundamental frequency. In addition, female bellows were significantly shorter in duration and had higher fundamental frequency, formant frequencies, and formant frequency spacing than male bellows. These results indicate that female koala vocalisations have the potential to signal the caller’s identity, age and sex. I go on to discuss the anatomical basis for these findings, and consider the possible functional relevance of signalling this type of information in the koala’s natural habitat.     

Bat Avoidance in Non-Aerial Insects: The Silence Response of Signaling Males in an Acoustic Moth

While the evasive responses of many flying acoustic insects to aerial‐hawking bats are duly recognized and studied, the responses of non‐aerial insects to gleaning bats are generally overlooked. It has been assumed that acoustic insects are deaf to these predators because gleaning bat echolocation calls are typically low in amplitude, brief (1–3 ms) and very high in frequency (>60 kHz). We tested this assumption in a series of playback experiments with a moth (Achroia grisella) that uses hearing in both predator evasion and mating. We report that ultrasound pulses ≥78 dB peSPL (peak equivalent sound pressure level) and ≥1 ms in duration inhibit stationary males from broadcasting their own ultrasonic advertisement calls, provided that the pulsed stimuli are delivered at a repetition rate ≤30/s. Further analyses suggest that inhibition by pulsed ultrasound comprises two processes performed serially. First, a startle response with a latency <50 ms is elicited by a single pulse ≥1 ms duration. Here, a male misses broadcasting several calls over a 50–100 ms interval. Secondly, the startle may be extended as a silence response lasting several to many seconds if subsequent pulses occur at a rate ≤30/s. Call inhibition cannot represent a simple response to acoustic power because of the inverse interaction between pulse duration and rate. On the other hand, the temporal and energy characteristics of inhibitory stimuli match those of gleaning bat echolocation calls, and we infer that inhibition is a specialized defensive behavior by which calling males may avoid detection by eavesdropping bats.     

Divergence of Acoustic Signals in a Widely Distributed Frog: Relevance of Inter-Male Interactions

Divergence of acoustic signals in a geographic scale results from diverse evolutionary forces acting in parallel and affecting directly inter-male vocal interactions among disjunct populations. Pleurodema thaul is a frog having an extensive latitudinal distribution in Chile along which males'' advertisement calls exhibit an important variation. Using the playback paradigm we studied the evoked vocal responses of males of three populations of P. thaul in Chile, from northern, central and southern distribution. In each population, males were stimulated with standard synthetic calls having the acoustic structure of local and foreign populations. Males of both northern and central populations displayed strong vocal responses when were confronted with the synthetic call of their own populations, giving weaker responses to the call of the southern population. The southern population gave stronger responses to calls of the northern population than to the local call. Furthermore, males in all populations were stimulated with synthetic calls for which the dominant frequency, pulse rate and modulation depth were varied parametrically. Individuals from the northern and central populations gave lower responses to a synthetic call devoid of amplitude modulation relative to stimuli containing modulation depths between 30–100%, whereas the southern population responded similarly to all stimuli in this series. Geographic variation in the evoked vocal responses of males of P. thaul underlines the importance of inter-male interactions in driving the divergence of the acoustic traits and contributes evidence for a role of intra-sexual selection in the evolution of the sound communication system of this anuran.     

Acoustic Signals of a Dog and Cat Induce Hemodynamic Responses within the Human Brain

The objective of this study was to evaluate the response of the human frontal cortex when listening to cat meows and dog barks, accompanied with a non-verbal pictorial using an affective rating system that assesses the dimensions of valence and arousal. Each participant (24 students; 12 females and 12 males) sat individually in the middle of a room and listened to the sounds (cat meows, dog barks, and the sound of a train) through ceiling-mounted speakers with a near infrared spectroscopic (NIRS) device. Participants had significantly higher oxygenated hemoglobin (oxy-Hb) levels during the exposure to meows (p < 0.05) and barks (p < 0.01) compared with the train sound. A significant correlation was observed between the dimensions of valence and oxy-Hb activation when the participants listened to cat meows (r = 0.53, p < 0.017; Bonferroni correction). In conclusion, we found that the acoustic signals of companion animals lead to frontal cortex responses in humans, suggesting that their signals have an important function related to their long coexistence with people.     

Responsiveness of cortisol and dehydroepiandrosterone to ACTH in children

In a total of 101 children, the dehydroepiandrosterone (DHA) and cortisol (F) levels were measured before and after ACTH (Synacten) administration. F responsiveness was unchanged during development, while DHA responsiveness in healthy children was highest during adrenarche. In hypopituitary patients DHA levels were lower than in the controls, but responsiveness to ACTH showed similar changes during development. Children with Turner's syndrome and hypergonadotrophic males had the response in elevated DHA levels while ACTH-induced DHA response related to bone-age matched controls. We conclude that regulation of adrenal androgens is mediated by both ACTH and another hypothalamo-pituitary hormone, perhaps independent of gonadal activation, but requiring gonadal integrity.     

Acoustic Signals of Katydids of the Tribe Gampsocleidini (Orthoptera,Tettigoniidae) in Russia

Entomological Review - Temporal patterns and frequency spectra of the acoustic calling signals of males of Uvarovites inflatus and four species of the genus Gampsocleis occurring in Russia are...