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  • 1
    Electronic Resource
    Electronic Resource
    Tympanal Hearing in Insects (1996)
    Palo Alto, Calif. : Annual Reviews
    Annual Review of Entomology 41 (1996), S. 433-450 
    Details
    ISSN: 0066-4170
    Source: Annual Reviews Electronic Back Volume Collection 1932-2001ff
    Topics: Biology
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1146/annurev.en.41.010196.002245
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  • 2
    Electronic Resource
    Electronic Resource
    Startle, Categorical Response, and Attention in Acoustic Behavior of Insects (1989)
    Palo Alto, Calif. : Annual Reviews
    Annual Review of Neuroscience 12 (1989), S. 355-375 
    Details
    ISSN: 0147-006X
    Source: Annual Reviews Electronic Back Volume Collection 1932-2001ff
    Topics: Biology , Medicine
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1146/annurev.ne.12.030189.002035
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  • 3
    Electronic Resource
    Electronic Resource
    Frequency as a releaser in the courtship song of two crickets, Gryllus bimaculatus (de Geer) and Teleogryllus oceanicus: a neuroethological analysis (1994)
    Springer
    Journal of comparative physiology 174 (1994), S. 485-494 
    Details
    ISSN: 1432-1351
    Keywords: Cricket ; Courtship ; Acoustic interneuron ; Neuroethology ; Playback
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract 1. The courtship behavior of male field crickets, Gryllus bimaculatus (De Geer) and Teleogryllus oceanicus, is a complex, multimodal behavioral act that involves acoustic signals (a courtship song; Fig. 1A,B). The dominant frequency is 4.5 kHz for T. oceanicus song (Fig. 1A) and 13.5 kHz for G. bimaculatus (Fig. IB). 2. When courting males are deprived of their courtship song by wing amputation, their courtship success declines markedly but is restored when courting is accompanied by tape-recordings of their courtship songs or a synthetic courtship song with only the dominant frequency of the natural song; other naturally occurring frequency components are ineffective for restoring mating success (Figs. 4, 5). 3. It has been suggested that an identified auditory interneuron, AN2, plays a critical role in courtship success. Chronic recordings of AN2 in an intact, tethered female show that AN2's response to the natural courtship song and synthesized songs at 4.5 and 13.5 kHz is similar in T. oceanicus. By contrast, in G. bimaculatus, AN2's response to the natural courtship song and synthesized song at 13.5 kHz, but not at 4.5 kHz, is similar (Figs. 2,3). 4. In behavioral experiments, playback of a 30 kHz synthetic courtship song in G. bimaculatus does not restore courtship success, yet this same stimulus elicits as strong a response from AN2 as does the normal courtship song (Fig. 6). Thus, contrary to earlier work by others, we conclude AN2 is not, by itself, a critical neural link in the courtship behavior of these two species of crickets.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/BF00191714
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  • 4
    Electronic Resource
    Electronic Resource
    Tympanal mechanics in the parasitoid fly Ormia ochracea : intertympanal coupling during mechanical vibration (1998)
    Springer
    Journal of comparative physiology 183 (1998), S. 443-452 
    Details
    ISSN: 1432-1351
    Keywords: Key words Directional hearing ; Tympanal ear Laser vibrometry ; Audition ; Auditory mechanics
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract The acoustic parasitoid fly Ormia ochracea locates its host, a singing field cricket, by means of a pair of small tympanal organs which are less than 2 mm in width. Nevertheless, laser vibrometric evidence shows that this tympanal system is directionally sensitive to sound through the action of a flexible intertympanal bridge that mechanically couples the tympana. Biomechanical data, a mechanical analogue and an analytical model lead to a testable prediction about the vibratory behavior of this tympanal system: if intertympanal coupling occurs, a force applied only unilaterally in non-acoustical conditions should be transmitted, at least to some degree, to the contralateral ear. This paper presents new experiments of direct mechanical stimulation that test the prediction of mechanical coupling. Stimulation on only one side of the intertympanal bridge elicited a contralateral mechanical response. Thus, coupling of the tympanal membranes through a flexible intertympanal bridge is demonstrated by mechanical as well as acoustical stimulation. These experiments also test for the possible presence of a pressure-difference system in O. ochracea. Intertympanal coupling is shown not to depend on the integrity of the air space backing the tympanal system, thus eliminating this possibility.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/s003590050270
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  • 5
    Electronic Resource
    Electronic Resource
    Peripheral frequency mis-match in the primitive ensiferan Cyphoderris monstrosa (Orthoptera: Haglidae) (1999)
    Springer
    Journal of comparative physiology 184 (1999), S. 543-551 
    Details
    ISSN: 1432-1351
    Keywords: Key words Auditory physiology ; Insect ; Frequency discrimination ; Evolution ; Song recognition
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract Peripheral auditory frequency tuning in the ensiferan insect Cyphoderris monstrosa (Orthoptera: Haglidae) was examined by comparing tympanal vibrations and primary auditory receptor responses. In this species there is a mis-match between the frequency of maximal auditory sensitivity and the frequency content of the species' acoustic signals. The mis-match is not a function of the mechanical properties of the tympanum, but is evident at the level of primary receptors. There are two classes of primary receptors: low-tuned and broadly tuned. Differences in the absolute sensitivity of the two receptor types at the male song frequency would allow the auditory system to discriminate intraspecific signals from sounds containing lower frequencies. Comparisons of tympanal and receptor tuning indicated that the sensitivity of the broadly tuned receptors did not differ from that of the tympanum, while low-tuned receptors had significantly narrower frequency tuning. The results suggest that the limited specialization for the encoding of intraspecific signals in the auditory system of C. monstrosa is a primitive rather than a degenerate condition. The limited specialization of C. monstrosa may reflect the evolutionary origin of communication-related hearing from a generalized precursor through the addition of peripheral adaptations (tympana, additional receptors) to enhance frequency sensitivity and discrimination.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/s003590050354
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  • 6
    Electronic Resource
    Electronic Resource
    Directional hearing by mechanical coupling in the parasitoid fly Ormia ochracea (1996)
    Springer
    Journal of comparative physiology 179 (1996), S. 29-44 
    Details
    ISSN: 1432-1351
    Keywords: Insect bioacoustics ; Sound localization ; Tympanal ear ; Laser vibrometry ; Ormia ochracea
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract Sound localization is a basic processing task of the auditory system. The directional detection of an incident sound impinging on the ears relies on two acoustic cues: interaural amplitude and interaural time differences. In small animals, with short interaural distances both amplitude and time cues can become very small, challenging the directional sensitivity of the auditory system. The ears of a parasitoid fly Ormia ochracea, are unusual in that both acoustic sensors are separated by only 520 μm and are contained within an undivided air-filled chamber. This anatomy results in minuscule differences in interaural time cues (ca. 2 μs) and no measurable difference in interaural intensity cues generated from an incident sound wave. The tympana of both ears are anatomically coupled by a cuticular bridge. This bridge also mechanically couples the tympanana, providing a basis for directional sensitivity. Using laser vibrometry, it is shown that the mechanical response of the tympanal membranes has a pronounced directional sensitivity. Interaural time and intensity differences in the mechanical response of the ears are significantly larger than those available in the acoustic field. The tympanal membranes vibrate with amplitude differences of about 12 dB and time differences on the order of 50 μs to sounds at 90° off the longitudinal body axis. The analysis of the deflection shapes of the tympanal vibrations shows that the interaural differences in the mechanical response are due to the dynamic properties of the tympanal system and reflect its intrinsic sensitivity to the direction of a sound source. Using probe microphones and extracellular recording techniques, we show that the primary auditory afferents encode sound direction with a time delay of about 300 μs. Our data point to a novel mechanism for directional hearing in O. ochracea based on intertympanal mechanical coupling, a process that amplifies small acoustic cues into interaural time and amplitude differences that can be reliably processed at the neural level. An intuitive description of the mechanism is proposed using a simple mechanical model in which the ears are coupled through a flexible lever.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/BF00193432
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  • 7
    Electronic Resource
    Electronic Resource
    The sounds of silence: cessation of singing and song pausing are ultrasound-induced acoustic startle behaviors in the katydid Neoconocephalus ensiger (Orthoptera; Tettigoniidae) (2000)
    Springer
    Journal of comparative physiology 186 (2000), S. 129-142 
    Details
    ISSN: 1432-1351
    Keywords: Key words Bushcricket hearing ; Central pattern generator ; Escape response ; Neural oscillator ; Predation
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract Previous studies of acoustic startle in insects have dealt with behavioral and/or neural mechanisms employed in evading aerially hawking, echolocating bats; however, insects also face terrestrial predators. Here we describe an acoustic startle response of the nocturnal katydid, Neoconocephalus ensiger. Stridulating males disturbed in the field perform obvious anti-predatory behaviors – cessation of singing, freezing, jumping, and evasive flight. Under controlled laboratory conditions we found that cessation of singing and song pausing are ultrasound-specific behaviors: when stimulated with pulsed ultrasound (20–100 kHz), but not audio-sound (〈20 kHz), males cease mate calling or insert pauses in their song. A second factor influencing acoustic startle is the phase of stimulation: an acoustic startle response occurs only when the pulse of ultrasound arrives during the window of silence between stridulatory syllables. The average startle threshold and response latency was 70 ± 5 dB SPL and 34.2 ± 6.0 ms, respectively. N. ensiger is particularly useful for examining acoustic startle responses of non-flying insects because (1) its calling song is broadband and contains ultrasound, thus the possibility exists of confusion over the biological meaning of ultrasound, and (2) this species shows the classic bat-avoidance response while flying, so a direct comparison between two types of acoustic startle is possible within the same species.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/s003590050013
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  • 8
    Electronic Resource
    Electronic Resource
    The response of an insect parasitoid, Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation (1995)
    Springer
    Behavioral ecology and sociobiology 36 (1995), S. 111-118 
    Details
    ISSN: 1432-0762
    Keywords: Key words Tachinid ; Uncertainty ; Optimal ; Parasitoid ; Clutch size
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Abstract  Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were superparasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. By manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7±1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1±5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.S.A Adamo ( ♪ ) ⋅ D. Robert ⋅ J. Perez ⋅ R.R. HoySection of Neurobiology and Behavior) S.G. Mudd Hall, Cornell University, Ithaca NY 14853–2702, USA
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/s002650050131
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  • 9
    Electronic Resource
    Electronic Resource
    The response of an insect parasitoid, Ormia ochracea (Tachinidae), to the uncertainty of larval success during infestation (1995)
    Springer
    Behavioral ecology and sociobiology 36 (1995), S. 111-118 
    Details
    ISSN: 1432-0762
    Keywords: Tachinid ; Uncertainty ; Optimal Parasitoid ; Clutch size
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology
    Notes: Abstract Ormia ochracea is a parasitoid fly which lays its larvae on its hosts, the field crickets Gryllus integer and Gryllus rubens, in two distinct modes: (1) directly on the host and (2) around the host. In the field, 12.7% of male crickets were parasitized and 3.2% were super-parasitized. Despite the disadvantages of parasitizing infested hosts, there was no evidence that O. ochracea avoided superparasitism. This and other experiments suggest that the host assessment ability of O. ochracea is less than that reported for many hymenopteran parasitoids. by manipulating the number of larvae in each cricket, we determined that four to five larvae per host resulted in the largest number of adult flies. However, as larval number per host increased from one to six, pupal size, and hence adult size, declined. In the field, hosts were found with a mean of 1.7 ± 1.0 (SD) larvae per cricket, suggesting that there may be some selection pressure against larger clutch sizes. Nevertheless clutch sizes larger than the host can support were sometimes found in the field. During the first mode of larviposition, gravid flies deposited no more than three larvae directly onto the host. Larvae deposited directly on the host had a high probability of infesting it. During the second mode of larviposition, gravid flies laid a larger number of larvae around the host (6.1 ± 5.2). Larvae that were laid around the host were less likely to infest a cricket than were larvae that were deposited directly onto it. O. ochracea is unique in that its two different modes of larviposition have different probabilities of larval success. Even though the success rate for larvae laid during the second mode of larviposition was low, the possibility of parasitizing more hosts appears to have selected for flies laying more larvae (e.g. increasing clutch size) than is optimal if all the larvae successfully entered a single host.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/BF00170716
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  • 10
    Electronic Resource
    Electronic Resource
    The tympanal hearing organ of the parasitoid fly Ormia ochracea (Diptera, Tachinidae, Ormiini) (1994)
    Springer
    Cell & tissue research 275 (1994), S. 63-78 
    Details
    ISSN: 1432-0878
    Keywords: Insect hearing ; Acoustic parasitoid ; Tympanal organ ; Scolopophorous mechanoreceptor ; Sexual dimorphism ; Convergent evolution ; Ormia ochracea (Insecta, Diptera)
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Medicine
    Notes: Abstract Tympanate hearing has evolved in at least 6 different orders of insects, but had not been reported until recently in the Diptera. This study presents a newly discovered tympanal hearing organ, in the parasitoid tachinid fly, Ormia ochracea. The hearing organ is described in terms of external and internal morphology, cellular organization of the sensory organ and preliminary neuroanatomy of the primary auditory afferents. The ear is located on the frontal face of the prothorax, directly behind the head capsule. Conspicuously visible are a pair of thin cuticular membranes specialized for audition, the prosternal tympanal membranes. Directly attached to these membranes, within the enlarged prosternal chamber, are a pair of auditory sensory organs, the bulbae acusticae. These sensory organs are unique among all auditory organs known so far because both are contained within an unpartitioned acoustic chamber. The prosternal chamber is connected to the outside by a pair of tracheae. The cellular anatomy of the fly's scolopophorous organ was investigated by light and electron microscopy. The bulba acustica is a typical chordotonal organ and it contains approximately 70 receptor cells. It is similar to other insect sensory organs associated with tympanal ears. The similarity of the cellular organization and tympanal morphology of the ormiine ear to the ears of other tympanate insects suggests that there are potent constraints in the design features of tympanal hearing organs, which must function to detect high frequency auditory signals over long distances. Each sensory organ is innervated by a branch of the frontal nerve of the fused thoracic ganglia. The primary auditory afferents project to each of the pro-, meso-, and metathoracic neuropils. The fly's hearing organ is sexually dimorphic, whereby the tympanal membranes are larger in females and the spiracles larger in males. The dimorphism presumably reflects differences in the acoustic behavior in the two sexes.
    Type of Medium: Electronic Resource
    URL: http://dx.doi.org/10.1007/BF00305376
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