Klaus Kalmring

Projection areas and branching patterns of the tympanal receptor cells in migratory locusts, Locusta migratoria and Schistocerca gregaria.

SummaryIn Locusta migratoria and Schistocerca gregaria, the projection areas and branching patterns of the tympanal receptor cells in the thoracic ganglia were revealed. Four auditory neuropiles can be distinguished on each side of the ventral cord, always located in the anterior part of the ring tract in each neuromere (two in the meta-, one in the meso-, and one in the prothoracic ganglion). Some of the receptor fibres ascend to the suboesophageal ganglion. There are distinct subdivisions within the auditory, frontal metathoracic and mesothoracic neuropiles. The arrangement of the terminal arborisations of the four types of tympanal receptor cells according to their different frequency-intensity responses is somatotopic and similar in the two ganglia. Here the receptor cells of type-1 form a restricted lateroventral arborisation. Cells of type-4 occupy the caudal part with a dorsorostral extension. Cells of type-2 and -3 arborise in a subdivision between both. Most of the stained low-frequency receptors (type-1, -2, and -3) terminate either in the metathoracic or, predominantly, in the mesothoracic ganglion. In contrast, the high-frequency cells (type-4) ascend to the prothoracic ganglion. The receptor fibres of the different types of receptor cells differ in diameter.

Behavioural adaptations of ground living bushcrickets to the properties of sound propagation in low grassland

SummaryThe acoustic behaviour of the closely related tetigoniid species Psorodonotus illyricus and Decticus verrucivorus have been invectigated by bioacoustical and behavioural methods. Both species show adaptations concerning the acoustic behaviour with respect to the biotope and the properties of sound propagation. These insects inhabit low grassland with an average vegetation height of about 20 cm which is also the general height for the song perches. Difficulties arise for efficient acoustic communication in such habitats. Sound propagation is influenced and limited by the strong ground attenuation and the excess damping by grass vegetation. Other limiting factors are the microclimatic conditions in the biotope. The two species counteract these difficulties by moving around in the biotope during stridulation. Both species mainly stridulate in the morning, avoiding problems of reduced sound transmission which often appear in the afternoon due to negative temperature gradients and resulting shadow zones. From the high mobility of these insects, it follows that individuals have no fixed territory and consequently no rivalry against conspecifics, which is very common among Orthopterans with a high degree of territoriality. It can be concluded that the preferred biotope influences and creates behavioural patterns in Orthopterans, especially here in the two investigated species of bushcrickets Psorodonotus illyricus and Decticus verrucivorus.

Structure of atympanate tibial organs in legs of the cave-living ensifera, Troglophilus neglectus (Gryllacridoidea, Raphidophoridae)

Troglophilus neglectus (Gryllacridoidea, Raphidophoridae) is a nocturnal Ensifera which can be found in caves of Slovenia. The anatomy of the tibial organs in the fore‐, mid‐, and hindlegs, as well as the external morphology of the proximal fore‐tibia and the prothoracic tracheal system, is described comparatively. In the prothorax and in the forelegs, no sound‐conducting structures such as an acoustic trachea, enlarged spiracles, or tympana are developed. A group of 8–10 campaniform sensillae is located in the dorsal cuticle of the proximal tibia. In each leg, the tibial organ complex is built up by two scolopale organs, the subgenual organ and the intermediate organ; the structure and the number of scolopidia is similar in each leg. No structure resembling the crista acoustica is found. The subgenual organ contains around 30 scolopidia; the intermediate organ is subdivided into a proximal part containing 8‐9 scolopidia and a distal part with 5–6 scolopidia. The two groups of scolopidia are not directly connected to the tracheal system. The tibial organs in the forelegs are insensitive to airborne sound, and they appear to be more primitive compared to those found in members of the Tettigoniidae and the Gwllidae. The results indicate that the complex tibial organs in all legs of T. neglectus are primarily vibrosensitive.

Functional morphology of bushcricket ears: comparison between two species belonging to the Phaneropterinae and Decticinae (Insecta, Ensifera)

SummaryThe morphology of the complex tibial organs in the forelegs of two bushcricket species belonging to the Phaneropterinae and Decticinae (Tettigoniidae) is described comparatively. In both species the tibial organs are made up of the subgenual organ, the intermediate organ and the crista acustica; the latter are parts of the tympanal organs and serve as auditory receptors. The very thin tympana in the forelegs ofPholidoptera griseoaptera (Decticinae) are protected by tympanal covers whereas inLeptophyes punctatissima (Phaneropterinae) the tympana are thicker and fully exposed. The overall auditory sensitivity ofL. punctatissima is lower and the sensitivity maximum of the hearing threshold lies at higher frequencies compared toP. griseoaptera. The number of scolopidia in the three scolopale organs and the dimensions of parts of the sound conducting system differs in the two species. In the crista acustica ofL. punctatissima a higher number of scolopidia is distributed in a smaller range than inP. griseoaptera; the scolopidia are especially concentrated in the distal part. Morphometrical analyses indicate that the dimensions of the spiracles, the acoustic trachea and the tympana determine the overall auditory sensitivity and that the arrangement of the scolopidia and the dimensions of structures in the crista acustica affect the frequency tuning of the hearing threshold.

Acoustic transmission characteristics of the tympanal tracheae of bushcrickets (Tettigoniidae). II: Comparative studies of the tracheae of seven species

The transmission characteristics of the acoustic tracheae in the forelegs of seven tettigoniid species were investigated by sinusoidal analysis. The species were selected to represent a range of body sizes and leg lengths. Four subfamilies were included, with two species each from three of them; the tracheae in such closely related pairs could be expected to be similar in shape despite their different dimensions. The tracheae were dissected out for morphometric analysis and compared with one another with respect to their overall dimensions and those of typical subsections. The amplitude‐versus‐frequency response of acoustic transmission in the tracheae was measured at various positions with a probe microphone. The stimuli were continuous sinusoidal signals at an intensity of 100 or 110 dB SPL. The tracheae of all the species studied here (in males and females) are distinguished by a bandpass‐limited transmission characteristic. In the frequency range above 5 kHz (at least to 40 kHz) the sound signals are amplified by 10–15 dB during passage through the tracheae. These results are compared with the threshold curves of the auditory organs and the spectra of the conspecific songs. Although in some cases there are considerable differences in the dimensions of the tracheae, the transmission characteristics are very similar; no specific adaptations to the frequency composition of the conspecific song were found.

Changes in the auditory system of locusts (Locusta migratoria and Schistocerca gregaria) after deafferentation

SummaryDeafferentation experiments during postembryonic development show morphological and/or physiological changes of receptor fibers and of identified auditory interneurons in the CNS of the locusts Locusta migratoria and Schistocerca gregaria after unilateral ablation of one tympanic organ either in the larva or the adult animal.1.In Locusta migratoria, 5 days after deafferentation, intact, contralateral receptor fibers had sprouted collaterals in the frontal acoustic neuropil of the metathoracic ganglion (Figs. 1, 2). Collateral sprouts were only rarely found in Schistocerca gregaria.2.After about 20 days the deafferented auditory interneurons receive new inputs from the contralateral receptors (Figs. 3, 5, 7, 10). This largely restores their thresholds and intensity/response functions. Collaterals from the first order interneurons cross the midline to the contralateral neuropil (BSN1 neuron, Fig. 4), which is never seen in intact animals. By contrast, in the TN1 neuron no consistent morphological change due to the deafferentation could be found (Fig. 6).3.Interneurons of higher order (AN1, TN3 neuron in locusts) regain their response pattern (Fig. 7) without morphological changes (Fig. 9). Bilateral recordings show that the deafferented interneurons respond more weakly to auditory stimuli than the intact neuron, but the response to vibration stimuli remains unchanged (TN3 neuron, Fig. 8).

The coding of airborne-sound and vibration signals in bimodal ventral-cord neurons of the grasshopperTettigonia cantans

SummaryIn grasshoppers, the auditory and vibrational senses converge on the same ventral-cord neurons. All neurons in the ventral cord that discharge impulses in response to either airborne-sound or vibration stimuli also receive synaptic inputs from the other sensory system. The latter elicit either subthreshold excitation or inhibition.The coding of the conspecific song in the responses of most ventral-cord neurons ofTettigonia cantans is considerably improved when the stimulus consists not of simulated natural sounds alone, but of such sounds together with either maintained vibration or vibration matched to the temporal structure of the song.Stridulating tettigoniids produce both airborne and substrate-conducted sound. Thus the perception of airborne sound and vibration, and their simultaneous processing in individual ventral-cord neurons, may be of fundamental importance — not only in localizing a nearby sound source, but also in facilitating the recognition of conspecific signals.

SOUND PRODUCTION AND SOUND EMISSION IN SEVEN SPECIES OF EUROPEAN TETTIGONIIDS. PART I. THE DIFFERENT PARAMETERS OF THE SONG; THEIR RELATION TO THE MORPHOLOGY OF THE BUSHCRICKET

Abstract Comparative studies of sound production and sound emission in seven species of European tettigoniids have been carried out. The species chosen were two Tettigoniines (Tettigonia cantans, Tettigonia viridissima), two Ephippigerines (Ephippiger discoidalis, Ephippiger ephippiger), and three Decticines (Decticus albifrons, Decticus verrucivorus, Psorodonotus illyricus). The factors which determined the choice of species were the different morphology (for example body shape and weight, and wing size) of the three subfamilies. The parameters of the different songs (e.g. dominant frequency, intensity) are normally not correlated to any of the investigated morphological characteristics of the animals. In the brachypterous species intraspecific correlations exist between wing size and the dominant low frequency band of the call. This frequency band is also observable at related higher frequencies in the ultrasonic range (20–60 kHz), the observed band width increasing with frequency. Sound emission in all...

Stimulus transmission in the auditory receptor organs of the foreleg of bushcrickets (Tettigoniidae) I. The role of the tympana.

The auditory organs of the tettigoniid are located just below the femoral tibial joint in the forelegs. Structurally each auditory organ consists of a tonotopically organized crista acustica and intermediate organ and associated sound conducting structures; an acoustic trachea and two lateral tympanic membranes located at the level of the receptor complex. The receptor cells and associated satellite structures are located in a channel filled with hemolymph fluid. The vibratory response characteristics of the tympanic membranes generated by sound stimulation over the frequency range 2-40 kHz have been studied using laser vibrometry. The acoustic trachea was found to be the principal structure through which sound energy reached the tympana. The velocity of propagation down the trachea was observed to be independent of the frequency and appreciably lower than the velocity of sound in free space. Structurally the tympana are found to be partially in contact with the air in the trachea and with the hemolymph in the channel containing the receptor cells. The two tympana were found to oscillate in phase, with a broad band frequency response, have linear coherent response characteristics and small time constant. Higher modes of vibration were not observed. Measurements of the pattern of vibration of the tympana showed that these structures vibrate as hinged flaps rather than vibrating stretched membranes. These findings, together with the morphology of the organ and physiological data from the receptor cells, suggest the possibility of an impedance matching function for the tympana in the transmission of acoustic energy to the receptor cells in the tettigoniid ear.

The auditory-vibratory system of the bushcricket Polysarcus denticauda (Phaneropterinae, Tettigoniidae) I. Morphology of the complex tibial organs

The structure of the complex tibial organs in the fore-, mid- and hindlegs of the bushcricket Polysarcus denticauda (Tettigoniidae, Phaneropterinae) is described comparatively. As is common for bushcrickets, in each leg the tibial organs consist of the subgenual and intermediate organs and the crista acustica. Only in the forelegs are sound-transmitting structures present. They consist of the spiracle, acoustic trachea, and two tympana; the latter are not protected by tympanal covers. The tympana in P. denticauda are extremely thick, not only bordering the two tracheal branches to the outside but also forming the outer wall of the hemolymph channel. The morphology of the tracheae in the mid- and hindlegs is significantly different, causing structural differences, especially in dimensions of the hemolymph channel. The number of scolopidia of the crista acustica of the foreleg is extremely high for a bushcricket. Approximately 50 receptor cells were found, about half of them being located in the distal quarter of the long axis of this organ. Some of the receptors are positioned in parallel on the dorsal wall of the anterior tracheal branch. The number, morphology and dimensions of the scolopidia within the crista acustica of the mid- and hindlegs differ significantly from those of the forelegs, decreasing in both legs to eight and seven receptor cells, respectively. Although the dimensions of the subgenual and intermediate organs are considerably larger in the mid- and hindlegs, the number of receptor cells is approximately the same in the different legs, being somewhat higher in both receptor organs than in those of many other bushcricket species studied previously.