@article {53499, title = {Innate releasing mechanisms and fixed action patterns: basic ethological concepts as drivers for neuroethological studies on acoustic communication in Orthoptera}, journal = {Journal of Comparative Physiology A}, year = {2019}, month = {Jul-01-2019}, abstract = {

This review addresses the history of neuroethological studies on acoustic communication in insects. One objective is to reveal how basic ethological concepts developed in the 1930s, such as innate releasing mechanisms and fixed action patterns, have influenced the experimental and theoretical approaches to studying acoustic communication systems in Orthopteran insects. The idea of innateness of behaviors has directly fostered the search for central pattern generators that govern the stridulation patterns of crickets, katydids or grasshoppers. A central question pervading 50 years of research is how the essential match between signal features and receiver characteristics has evolved and is maintained during evolution. As in other disciplines, the tight interplay between technological developments and experimental and theoretical advances becomes evident throughout this review. While early neuroethological studies focused primarily on proximate questions such as the implementation of feature detectors or central pattern generators, later the interest shifted more towards ultimate questions. Orthoptera offer the advantage that both proximate and ultimate questions can be tackled in the same system. An important advance was the transition from laboratory studies under well-defined acoustic conditions to field studies that allowed to measure costs and benefits of acoustic signaling as well as constraints on song evolution.

}, keywords = {acoustic communication, Central pattern generators, Coevolution, Genetic coupling, Sender{\textendash}receiver evolution}, issn = {0340-7594}, doi = {10.1007/s00359-018-01311-3}, url = {http://link.springer.com/10.1007/s00359-018-01311-3}, author = {Bernhard Ronacher} } @article {53391, title = {The impact of age and egg-laying cycle on female grasshoppers{\textquoteright} preference functions for acoustic signals}, journal = {Journal of Comparative Physiology A}, year = {2018}, month = {Jan-12-2018}, abstract = {

Female responsiveness and the shape of preference functions for male signal traits are important determinants for male mating success. We observed the responsiveness and the selectivity of virgin grasshopper females (Chorthippus biguttulus L.) for different features of males\’ acoustic signals throughout their life span to detect possible influences of age on the females\’ preference functions. In particular, we explored the hypothesis that the females may become less selective with increasing age and, therefore, would start to accept songs that are normally rejected. Such an age effect could relieve the selection pressure on male signal traits. In the majority of animals tested, the general responsiveness decreased with age although a few individuals exhibited an opposite trend. Contrary to the above expectation, there was no indication of a loss of selectivity in older females or an increased acceptance of normally unattractive song models. The timing within the oviposition cycle had a strong effect on responsiveness: near oviposition the general responsiveness increased and with it also the half width of the preference functions. However, highly unattractive song models remained unattractive also near oviposition.

}, keywords = {acoustic communication, Age effects, Female choice, Oviposition, Preference functions}, issn = {0340-7594}, doi = {10.1007/s00359-018-1307-8}, url = {http://link.springer.com/10.1007/s00359-018-1307-8}, author = {Aufderheide, Jennifer and Bernhard Ronacher} } @article {53154, title = {Evolutionarily conserved coding properties favour the neuronal representation of heterospecific signals of a sympatric katydid species}, journal = {Journal of Comparative Physiology A}, volume = {204}, year = {2018}, month = {Jan-10-2018}, pages = {859 - 872}, abstract = {

To function as a mechanism in premating isolation, the divergent and species-specific calling songs of acoustic insects must be reliably processed by the afferent auditory pathway of receivers. Here, we analysed the responses of interneurons in a katydid species that uses long-lasting acoustic trills and compared these with previously reported data for homologous interneurons of a sympatric species that uses short chirps as acoustic signals. Some interneurons of the trilling species respond exclusively to the heterospecific chirp due to selective, low-frequency tuning and \“novelty detection\”. These properties have been considered as evolutionary adaptations in the sensory system of the chirper, which allow it to detect signals effectively during the simultaneous calling of the sympatric sibling species. We propose that these two mechanisms, shared by the interneurons of both species, did not evolve in the chirper to guarantee its ability to detect the chirp under masking conditions. Instead we suggest that chirpers evolved an additional, 2-kHz component in their song and exploited pre-existing neuronal properties for detecting their song under masking noise. The failure of some interneurons to respond to the conspecific song in trillers does not prevent intraspecific communication, as other interneurons respond to the trill.

}, keywords = {Auditory processing, insects, Selective coding, signal detection, signal evolution}, issn = {0340-7594}, doi = {10.1007/s00359-018-1282-0}, url = {http://link.springer.com/10.1007/s00359-018-1282-0}, author = {Kostarakos, Konstantinos} } @article {53148, title = {Song harmonic content as a parameter determining acoustic orientation behaviour in the cricketTeleogryllus oceanicus (Le Guillou)}, journal = {Journal of Comparative Physiology A}, volume = {158}, year = {1986}, pages = {583{\textendash}591}, abstract = {

Analyses of natural songs from the cricket Teleogryllus oceanicus confirm (after Hutchings and Lewis 1984) that the songs are rich in harmonics which extend up to around 55 kHz. A series of synthetic song models with a varying harmonic content were used to determine the relevance of these harmonics to the orientation behaviour of the insects.

In two-choice experiments in which song models were presented simultaneously, crickets showed a clear discrimination between a song with harmonics and a song without. Female insects orientated preferentially to a song with harmonics. Interestingly, males preferred a song model without harmonics in the two-choice situation.

Two experimental regimes were used to test orientation accuracy, a forced-choice (Y maze) phonotactic experiment and a study of free phonotaxis in the behavioural arena. The results of both experimental approaches confirm that crickets orientate more accurately to a song with harmonics and can discriminate smaller angles to sound sources ahead and lateral to the insect.

Presentations of song models in which the fundamental (5 kHz) was attenuated relative to the 2nd harmonic have given results that imply a process of neural integration (two-tone or side band suppression) of the fundamental and the high-frequency harmonics.

}, keywords = {Arena, Orientation Accuracy, Orientation Behaviour, Side Band, Small Angle}, author = {William Latimer and Lewis, DB} } @article {53146, title = {Auditory behavior of the cricket IV. Interaction of direction of tracking with perceived temporal pattern in split-song paradigms}, journal = {Journal of Comparative Physiology A}, volume = {163}, year = {1988}, pages = {13-22}, chapter = {13}, abstract = {

1. Split songs \— stimulus arrangements in which parts of cricket calling songs are delivered from different directions via two loudspeakers \— have been used to place constraints on the recognition and direction-determining processes in the field cricketGryllus bimaculatus, walking on the Kramer spherical treadmill. Parts of a calling song, each ineffective when delivered alone, can combine to elicit recognition and tracking when interleaved from two speakers 135\° apart in azimuth.

2. If two correct songs (40-ms syllable repetition interval) are delivered from the two speakers, but time-shifted by a half interval so that their composite (20-ms intervals) is incorrect, then the angle bisecting the speaker separation (where the two should appear equally loud) might be expected to be an \‘unfavored zone\’; the animals do in fact tend to avoid that region. As in controls on \‘choice\’ experiments with interleaving of complete chirps, if one speaker is increased in sound intensity it tends to be favored.

3. If two incorrect songs (80-ms syllable repetition interval) are delivered from the two speakers, again time-shifted by half a syllable interval so that the composite is correct (40-ms intervals), then angles near those bisecting the speaker separation might be expected to be favored, and most animals track in that way. Surprisingly, however, when one speaker is increased in sound intensity in this case most animals show a marked shift of tracking toward the less-loud speaker at test intensities near 60 and 70 dB. At 80 dB the effect is reversed.

4. About one-third of females track trill \— continuous trains of syllables at the correct syllable rate but with no subdivision into chirps; these were tested with split trills. If alternate syllables of the trill were delivered from the two speakers, or alternating doublets, tracking tended to bisect the 135\° angle between speakers. With alternating triplets or higher n-tuples, one or the other speaker was tracked quite accurately. This ability to lock on to one signal despite the presence of another (at 135\°) was tested at smaller speaker separations (90\°, 65\°, 45\°); the animals then usually fail to select one speaker.

5. The above results are difficult to explain via a putative \‘tracking comparator\’ with \‘cardioid\’ inputs from the two sides of the auditory system. In particular, the distinct behaviors with respect to intensity imbalance (2 and 3 above, for different ranges of syllable repetition interval of split songs) suggest that bilateral dynamic interactions in the CNS may be involved. Specific correlates of these effects can be sought in multiple-source or dual leg-phone experiments with electrophysiological monitoring of prothoracic and brain cells.

}, author = {Theo Weber and John Thorson} } @article {53135, title = {Insect hearing in the field II. Male spacing behaviour and correlated acoustic cues in the bushcricket Mygalopsis marki}, journal = {Journal of Comparative Physiology A}, volume = {159}, year = {1986}, pages = {627{\textendash}638}, abstract = {

1. Distances between calling maleMygalopsis marki Bailey (Orthoptera: Tettigoniidae) were measured in two geographically distinct populations. Mean intermale distances differed significantly in the two habitats (6.2 m and 11.5 m), but the sound pressure level of the nearest neighbour, measured at each focal male, was close to 65 dB SPL in both populations (Figs. 4, 6).

2. Using the response of an identified auditory interneuron as a \‘biological microphone\’, the maximum hearing distance of males was tested. It was found to be affected by the size of the calling male, its posture, perch height and the vegetation density. Perch height alone influenced the maximum broadcast area of males by some 100\%. Nearest neighbour distances were less than half the maximum hearing distances, confirming by neurophysiological methods that males do not space on threshold sensitivity to the complete call (Fig. 5).

3. The ability of the bushcricket to perceive the temporal characteristic of the call was also tested with the auditory interneuron over increasing distances from a signaller. The temporal integrity of the song was found to be maintained over the whole broadcasting area (Fig. 9).

4. The call has two energy peaks at 10 kHz and 20 kHz. The intensity of each peak attenuated differentially with increasing distance from the signaller; the low frequency component had a low variance at the nearest neighbour distance (SD: +4 dB/\−6 dB) compared to the high frequency component (SD: +5 dB/\−12.5 dB) (Fig. 8). The importance of the total sound pressure level and the relative intensity of the major frequency components of the song are discussed in relation to likely cues available to the male for maintaining spacing.

}, keywords = {Focal Male, Frequency Component, High Frequency Component, Neighbour Distance, Sound Pressure Level}, author = {Heiner R{\"o}mer and Winston J. Bailey} } @article {53089, title = {What does a butterfly hear? Physiological characterization of auditory afferents in Morpho peleides (Nymphalidae)}, journal = {Journal of Comparative Physiology A}, year = {2018}, month = {May-08-2020}, abstract = {

Many Nymphalidae butterflies possess ears, but little is known about their hearing. The tympanal membrane of butterflies typically comprises distinct inner and outer regions innervated by auditory nerve branches NII and NIII and their respective sensory organs. Using the Blue Morpho butterfly (Morpho peleides) as a model, we characterized threshold and suprathreshold responses of NII and NIII. Both are broadly tuned to 1\–20 kHz with best frequencies at 1\–3 kHz, but NIII is significantly more sensitive than NII. The compound action potentials (CAPs) of both branches increase their first peak amplitudes and areas in response to higher sound levels. NII and NIII differed in their suprathreshold CAP responses to sound frequencies, with stronger responses to 1\–3 and 4\–6 kHz, for NIII and NII respectively; results that are consistent with tympanal membrane mechanics. These results indicate that butterflies are capable of amplitude and frequency discrimination. Both auditory branches responded to playbacks of the flight and calls of predatory birds. We propose that the ears of butterflies, like those of many vertebrate prey such as some rabbits and lizards, function primarily in predator risk assessment.

}, keywords = {Butterfly, Ear, neurophysiology, Predator detection, Sensory}, issn = {0340-7594}, doi = {10.1007/s00359-018-1280-2}, url = {http://link.springer.com/10.1007/s00359-018-1280-2}, author = {Mikhail, Andrew and Lewis, John E. and Yack, Jayne E.} } @article {47308, title = {Long-distance, low-frequency elephant communication}, journal = {Journal of Comparative Physiology A}, volume = {190}, year = {2004}, month = {Jan-10-2004}, pages = {791 - 805}, abstract = {

The production, transmission, and reception of and the behavioral response to long-distance, low-frequency sound by elephants is reviewed. The structure of low-frequency calls generated by elephants is separated into the \“source\” and the \“filter\” roles played by the lungs, larynx and vocal track, the composition of the expired air and the ambient air temperature. Implications regarding the size, age, sex, sexual and physical status follow from the call structure and detection. Reception of the signal is discussed in terms of the characteristics of the elephant\’s ear with particular attention to the determination of the threshold of hearing and the ability to locate the source of low-frequency sounds. Factors which influence the transmission of near infrasound are related to atmospheric structure. The critical role played by the thermal stratification and vertical gradient and magnitude of the wind in determining both the range and the detection of a signal are discussed for open and closed elephant habitats. Infrasound plays a pervasive role in reproduction, resource utilization, avoidance of predation and other social interactions. Current and future technology can be expected to contribute to the detection and interpretation of elephant communication. This will aid in the understanding of behavior and in efforts to sustain the species.

}, keywords = {behavioral response, calling, Elephant communication, hearing, transmission}, issn = {0340-7594}, doi = {10.1007/s00359-004-0553-0}, url = {http://link.springer.com/10.1007/s00359-004-0553-0}, author = {Garstang, Michael} } @article {47297, title = {Auditory processing in the black-sided meadow katydid Conocephalus nigropleurum (Orthoptera: Tettigoniidae)}, journal = {Journal of Comparative Physiology A}, volume = {164}, year = {1989}, month = {Jan-01-1989}, pages = {501 - 512}, abstract = {

1. The auditory system of the tettigoniid,Conocephalus nigropleurum was examined using whole auditory nerve responses (averaged summed action potentials (SAPs)) and intracellularly recorded single cells (receptor and interneurone) to pure tone stimuli and con- and heterospecific (C. brevipennis) songs.

2.The morphology of the auditory tracheal system and crista acustica is documented forC. nigropleurum. A broadly open thoracic spiracle (stigma) leads, via a horn, camera and foreleg trachea to a crista acustica of 28 receptor cells.

3. C. nigropleurum\&$\#$39;s auditory sensitivity is broadly tuned to 15\–35 kHz which is below the peak frequency band of its calling song (30\–36 kHz). The ear possesses a dynamic range of at least 30 dB and pulsed sounds with short (\<0.5 ms) onset times or long interpulse periods (\>200 ms) elicit maximal responses. We suggest this is caused by increased auditory receptor firing synchrony to these types of sounds and may facilitate the location of such sounds. There is, however, discrepancy between the pulse periods that elicit maximal SAP amplitudes and those found within the normal calling song, a phenomenon which may be related to the chorusing behaviour of this species.

4.Polar directional plots indicate thatC. nigropleurum is most sensitive to sounds originating posterior to the opening of the prothoracic auditory stigma; this may be caused by the configuration of the prothoracic pronotum. We found no pronounced sensitivity at points facing the tympanal slits.

5.Intracellular recordings reveal prothoracic receptors with widely decussate and narrowly branched termini. No receptor tested was able to track the intra-phonatome pulses (tooth strike sounds) of either species and we conclude that the phonatome is the finest resoluble component of the song.

6. An acoustically-activated interneurone (101) was recorded that resembles both the gryllid int-2 (Omega) cell and that of another tettigoniid. We suggest that tonic interneuronal firing caused by the multiple singer chorused song of either species activates the initial stages of phonotaxis in receptiveC. nigropleurum females but individual species recognition may depend at least partially on the discrimination of phonatome rates.

}, issn = {0340-7594}, doi = {10.1007/BF00610444}, url = {http://link.springer.com/10.1007/BF00610444}, author = {Fullard, James H. and Glenn K. Morris and Mason, Andrew C.} } @article {17230, title = {Separate localization of sound recognizing and sound producing neural mechanisms in a grasshopper}, journal = {Journal of Comparative Physiology A}, volume = {161}, year = {1987}, month = {Jan-01-1987}, pages = {95 - 101}, abstract = {

In the two acridid species Chorthippus parallelus and Ch. montanus, the sound template by which females recognize male song varies with temperature, as does the song itself. At relatively high temperatures the females respond best to simulated songs with high syllable frequencies, and at lower temperatures songs with lower syllable frequencies are preferred.

The temperature around the supraesophageal and metathoracic ganglia of female grasshoppers was monitored by implanted thermocouples, and either the head or the thorax was warmed selectively while the animal was free to move (within the imits of the wires). Then simulations of the conspecific song varying in syllable frequency corresponding to different song temperatures were presented, and the stridulatory responses of the animals were observed.

The results were as follows. 1. Song recognition (in particular, the position of the peak of the response curve) depended on the temperature of the head. 2. The rate of stridulatory hindleg movement was determined by the temperature of the thoracic ganglia.

This result provides strong evidence against the genetic coupling hypothesis.

}, issn = {0340-7594}, doi = {10.1007/BF00609458}, url = {http://link.springer.com/10.1007/BF00609458}, author = {Bauer, Maria and Otto von Helversen} } @article {17175, title = {Influence of syllable period on song encoding properties of an ascending auditory interneuron in the cricketAcheta domestica}, journal = {Journal of Comparative Physiology A}, volume = {165}, year = {1989}, month = {Jan-01-1989}, pages = {827 - 836}, issn = {0340-7594}, doi = {10.1007/BF00610881}, url = {http://link.springer.com/10.1007/BF00610881}, author = {Atkins, Steven and Atkins, Gordon and Rhodes, Mike and Stout, John F.} } @article {17174, title = {Response properties of prothoracic, interganglionic, sound-activated interneurons in the cricketTeleogryllus oceanicus}, journal = {Journal of Comparative Physiology A}, volume = {161}, year = {1987}, month = {Jan-01-1987}, pages = {681 - 693}, issn = {0340-7594}, doi = {10.1007/BF00605009}, url = {http://link.springer.com/10.1007/BF00605009}, author = {Atkins, Gordon and Gerald S. Pollack} } @article {17173, title = {Changes in phonotaxis by the female cricket Acheta domesticus L. after killing identified acoustic interneurons}, journal = {Journal of Comparative Physiology A}, volume = {154}, year = {1984}, month = {Jan-01-1984}, pages = {795 - 804}, issn = {0340-7594}, doi = {10.1007/BF00610680}, url = {http://link.springer.com/10.1007/BF00610680}, author = {Atkins, Gordon and Ligman, Scott and Burghardt, Frederick and Stout, John F.} } @article {12665, title = {Selective phonotaxis of female crickets under natural outdoor conditions}, journal = {Journal of Comparative Physiology A}, volume = {200}, year = {2014}, month = {Jan-03-2014}, pages = {239 - 250}, abstract = {Acoustic mate choice in insects has been extensively studied under laboratory conditions, using different behavioural paradigms. Ideally, however, mate choice designs should reflect natural conditions, including the physical properties of the transmission channel for the signal. Since little is known about the discrimination ability of females between male song variants under natural conditions, we performed phonotaxis experiments with female field crickets (Gryllus bimaculatus) outdoors, using two-choice decisions based on differences in carrier frequency, sound pressure level, and chirp rate. For all three song parameters, minimum differences necessary for a significant preference between two song models were considerably larger outdoors compared to laboratory conditions. A minimum amplitude difference of 5 dB was required for a significant choice in the field, compared to only 1{\textendash}2 dB reported for lab-based experiments. Due to the tuned receiver system, differences in carrier frequency equal differences in perceived loudness, and the results on choice for differences in carrier frequency corroborate those in amplitude. Similarly, chirp rate differences of 50 chirps/min were required outdoors compared to only 20 chirps/min in the lab. For predictions about patterns of sexual selection, future studies need to consider the different outcomes of mate choice decisions in lab and field trials.}, keywords = {acoustic communication, Decision making, Female choice, Field cricket, phonotaxis}, issn = {0340-7594}, doi = {10.1007/s00359-014-0881-7}, url = {http://link.springer.com/10.1007/s00359-014-0881-7}, author = {Hirtenlehner, Stefan} }