Neurophysiology of the vibration sense in locusts and bushcrickets: The responses of ventral-cord neurones

In the locustid Locusta migratoria and the tettigoniids Decticus verrucivorus and Tettigonia cantans, comparative aspects of physiological properties of vibratory/auditory ventral-cord neurones were studied by single cell recordings.These neurones all receive inputs from both vibratory and auditory receptors. Nevertheless, they can be classified into “V neurones” responding preferentially to vibration stimuli, “VS neurones” responding to vibration and airborne sound, and “S neurones” responding preferentially to airborne sound. In every group, there are several types with different physiological properties, normally represented by one neurone on each body side.In Locusta and in the tettigoniid species, the same physiological types of vibratory/auditory neurones were found, although there are differences in the synaptic connectivity of the vibration receptors of the different legs. In Locusta, the middle leg receptors have the strongest influence on the generation of suprathreshold responses of the central neurones, whereas in the tettigoniids the receptors of the ipsilateral fore leg are the most influential.Two of the V neurones receive inputs mainly from campaniform sensilla and other low-frequency vibration receptors, the other V and VS neurones are mainly influenced by the subgenual receptors. Central information processing results in preferential responses to different frequency/intensity ranges in different neurones.Most VS neurone types show the same response characteristics (e.g. time pattern of response, habituation) either to vibration or to airborne-sound stimuli. Simultaneous presentation of both stimuli leads to qualitative changes in the response characteristics. Therefore, the co-processing of auditory and vibratory signals seems to be very important in the acoustic behaviour of grasshoppers.     

Structure and sensory physiology of the leg scolopidial organs in Mantophasmatodea and their role in vibrational communication

Individuals of the insect order Mantophasmatodea use species-specific substrate vibration signals for mate recognition and location. In insects, substrate vibration is detected by mechanoreceptors in the legs, the scolopidial organs. In this study we give a first detailed overview of the structure, sensory sensitivity, and function of the leg scolopidial organs in two species of Mantophasmatodea and discuss their significance for vibrational communication. The structure and number of the organs are documented using light microscopy, SEM, and x-ray microtomography. Five scolopidial organs were found in each leg of male and female Mantophasmatodea: a femoral chordotonal organ, subgenual organ, tibial distal organ, tibio-tarsal scolopidial organ, and tarso-pretarsal scolopidial organ. The femoral chordotonal organ, consisting of two separate scoloparia, corresponds anatomically to the organ of a stonefly (Nemoura variegata) while the subgenual organ complex resembles the very sensitive organs of the cockroach Periplatena americana (Blattodea). Extracellular recordings from the leg nerve revealed that the leg scolopidial organs of Mantophasmatodea are very sensitive vibration receptors, especially for low-frequency vibrations. The dominant frequencies of the vibratory communication signals of Mantophasmatodea, acquired from an individual drumming on eight different substrates, fall in the frequency range where the scolopidial organs are most sensitive.     

Processing of vibratory and acoustic signals by ventral cord neurones in the cricket Gryllus campestris

The responses of single vibratory receptors and ascending ventral cord interneurones were studied extracellularly in Gryllus campestris L. The physiology of the vibration receptors resembled those found in tettigoniids and locusts. The frequency responses of the subgenual receptors provide two possible cues for central frequency discrimination: differences in mean tuning between groups of receptors in the different leg pairs and a range of receptors tuned to different frequencies within one subgenual organ.Most of the ascending vibratory interneurones were highly sensitive in either the low or high frequency range. Broadbanded neurones were less sensitive. The characteristic sensitivity peaks of these units are due mainly to receptor inputs from a particular leg pair, although most central neurones receive inputs from all 6 legs. Only one neurone type, TN1 received excitatory inputs from both auditory and vibratory receptors; its responses were greatly enhanced by the simultaneous presentation of both stimulus modes. The responses to sound stimuli of AN2, on the other hand, were inhibited by vibration. No other auditory interneurones investigated were influenced by inputs from vibration receptors. Central processing of vibratory information in the cricket is compared with that of tettigoniids and locusts.     

The responses of auditory ventral-cord neurons ofLocusta migratoria to vibration stimuli

Summary Most of the auditory neurons in the ventral nerve cord ofLocusta migratoria carry information not only from the tympanal organs but also from the subgenual organs (vibration sensors). Six of the eight neuron types studied electrophysiologically respond to at least these two modalities. Artificial sounds (white noise and pure tones varying in frequency and intensity) and sinusoidal vibration (200 Hz with an acceleration of 15.8 cm/s₂ or 2000 Hz and 87 cm/s₂) were used as stimuli.Complex excitatory and/or inhibitory interactions of the signals from both tympanal organs form the discharge patterns of auditory ventral-cord neurons in response to stimulation with air-borne sound. Normally the input of the ipsilateral sense organ dominates. The response patterns of these same neurons elicited by vibration stimuli are formed differently, as follows: (1) the sensory inputs of all subgenual organs are integrated in the responses of the ventral-cord neurons; in a single neuron they have either excitatory or inhibitory effects, but not both. (2) The more legs vibrated, the larger is the response. (3) The subgenual organs in the middle legs are most effective, those in the hind legs least so. (4) Ipsilateral vibration has more effect than contralateral.The six auditory neurons react to vibration combined with air-borne sound in different ways. The B neuron is the only one inhibited by vibration stimuli. The G neuron has been studied more intensively; because its anatomical arrangement and the location of the endings of the subgenual receptor fibers are known, it could be inferred from effects of transection of the connectives that interneurons are interposed between receptor cells and the G neuron.Part of the program Sonderforschungsbereich 114 (Bionach) Bochum, under the auspices of the Deutsche Forschungsgemeinschaft, with the support of the Slovenic Research Society (RSS)     

Functional morphology and development of tibial organs in the legs I,II and III of the bushcricketEphippiger ephippiger (Insecta,Ensifera)

Summary The anatomy of the complex tibial organs in the pro-, meso- and metathoracic legs of adults and larvae of the bushcricketEphippiger ephippiger is described comparatively. The subgenual organ and the intermediate organ are differentiated in the same way in legs I, II and III; the anatomy of the crista acustica and the tracheal morphology are significantly different. The final number of scolopidia in the tibial organ of each leg is present at the time of hatching. In the subgenual organ, the number of scolopidia is the same in all legs; in the intermediate organ, and especially in the crista acustica, the number of scolopidia decreases from leg I to legs II and III. In the first larval instar, the morphology of the tibia, the course of the trachea and the anatomy of accessory structures are developed in the same way in each leg. The specific differentiations forming the auditory receptor organ in leg I, such as the acoustic trachea, the tympana and tympanal cavities, develop step by step in subsequent instars. The auditory threshold recorded from the tympanal nerve in the prothoracic leg of adults is remarkably lower than in the meso- and metathoracic legs. Morphometrical analyses of structures that are suggested to play a role in stimulus transduction on scolopidia of the crista acustica reveal significant differences in the three legs.     

Biophysics of the subgenual organ of the honeybee, Apis mellifera

The subgenual organ of the honeybee (Apis mellifera) is suspended in a haemolymph channel in the tibia of each leg. When the leg is accelerated, inertia causes the haemolymph (and the subgenual organ) to lag behind the movement of the rest of the leg. The magnitude of this phase lag determines the displacement of the subgenual organ relative to the leg and to the proximal end of the organ, which is connected to the cuticle. Oscillations of the subgenual organ are visualised during vibration stimulation of the leg, by means of stroboscopic light. Video analysis provides fairly accurate values of the amplitude and phase of the oscillations, which are compared with the predictions of a model.   The model comparison shows that the haemolymph channel can be described as an oscillating fluid-filled tube occluded by an elastic structure (probably the subgenual organ). The mechanical properties of the subgenual organ and haemolymph channel resemble those of an overdamped mass-spring system. A comparison of the threshold curve of the subgenual organ determined using electrophysiology with that predicted by the oscillating tube model suggests that the sensory cells respond to displacements of the organ relative to the leg. Accepted: 10 May 1997     

Vibratory communication in spiders

We analyzed the response of the vibration sensitive lyriform organ on the metatarsus of female spiders (Cupiennius salei) to dummies of male courtship vibrations. One of the two representative slits studied is sharply tuned to 500 Hz. Only the other slit is sensitive enough at lower frequencies to represent the parameters contained in the behaviourally effective dummies:

1. Amplitude. The physiological threshold is similar to the behavioural threshold. The stimulus acceleration amplitudes leading to a good synchronization between response and temporal stimulus pattern coincide with those effectively eliciting a behavioural response. The most frequent spike intervals remain nearly constant in this range. At acceleration amplitudes above the natural range, syllable and pause durations are misrepresented by the receptor response.
2. Frequency. Varying the carrier frequency between 35–500 Hz changes the most frequent spike intervals. Interval histograms resulting from behaviourally effective stimuli (50–200 Hz), however, are similr for carrier frequencies differing by a factor of 2.
3. Temporal pattern. Response duration reflects the temporal parameters of the stimulus. The most frequent spike interval only changes with temporal stimulus characteristics far off the natural range. The number of spikes during a syllable decreases in ongoing stimulus series. The quality of copying the temporal stimulus pattern remains unchanged, however.

     

Human postural responses to different frequency vibrations of lower leg muscles

We analyzed human postural responses to muscle vibration applied at four different frequencies to lower leg muscles, the lateral gastrocnemius (GA) or tibialis anterior (TA) muscles. The muscle vibrations induced changes in postural orientation characterized by the center of pressure (CoP) on the force platform surface on which the subjects were standing. Unilateral vibratory stimulation of TA induced body leaning forward and in the direction of the stimulated leg. Unilateral vibration of GA muscles induced body tilting backwards and in the opposite direction of the stimulated leg. The time course of postural responses was similar and started within 1 s after the onset of vibration by a gradual body tilt. When a new slope of the body position was reached, oscillations of body alignment occurred. When the vibrations were discontinued, this was followed by rapid recovery of the initial body position. The relationship between the magnitude of the postural response and frequency of vibration differed between TA and GA. While the magnitude of postural responses to TA vibration increased approximately linearly in the 60-100 Hz range of vibration frequency, the magnitude of response to GA vibration increased linearly only at lower frequencies of 40-60 Hz. The direction of body tilt induced by muscle vibration did not depend on the vibration frequency.     

Vibration induced osteogenic commitment of mesenchymal stem cells is enhanced by cytoskeletal remodeling but not fluid shear

Consistent across studies in humans, animals and cells, the application of vibrations can be anabolic and/or anti-catabolic to bone. The physical mechanisms modulating the vibration-induced response have not been identified. Recently, we developed an in vitro model in which candidate parameters including acceleration magnitude and fluid shear can be controlled independently during vibrations. Here, we hypothesized that vibration induced fluid shear does not modulate mesenchymal stem cell (MSC) proliferation and mineralization and that cell's sensitivity to vibrations can be promoted via actin stress fiber formation. Adipose derived human MSCs were subjected to vibration frequencies and acceleration magnitudes that induced fluid shear stress ranging from 0.04 Pa to 5 Pa. Vibrations were applied at magnitudes of 0.15g, 1g, and 2g using frequencies of both 100 Hz and 30 Hz. After 14 d and under low fluid shear conditions associated with 100 Hz oscillations, mineralization was greater in all vibrated groups than in controls. Greater levels of fluid shear produced by 30 Hz vibrations enhanced mineralization only in the 2g group. Over 3 d, vibrations led to the greatest increase in total cell number with the frequency/acceleration combination that induced the smallest level of fluid shear. Acute experiments showed that actin remodeling was necessary for early mechanical up-regulation of RUNX-2 mRNA levels. During osteogenic differentiation, mechanically induced up-regulation of actin remodeling genes including Wiskott–Aldrich syndrome (WAS) protein, a critical regulator of Arp2/3 complex, was related to the magnitude of the applied acceleration but not to fluid shear. These data demonstrate that fluid shear does not regulate vibration induced proliferation and mineralization and that cytoskeletal remodeling activity may play a role in MSC mechanosensitivity.     

Anatomical and physiological observations on the organization of mechanoreceptors and local interneurons in the central nervous system of the wandering spider Cupiennius salei

Summary In the wandering spider Cupiennius salei, the functional neuroanatomy of leg mechanosensory receptor neurons and interneurons associated with a single leg neumere was investigated by combined intracellular recording and Lucifer yellow ionophoresis. Trichobothria axons that selectively respond to air currents and to low-frequency airborne vibrations have arborizations restricted to ventral regions of the appropriate leg neuromere. Receptor afferents that respond selectively to substrateborne vibrations are distributed ventrally in the corresponding leg neuromere and extend into certain interganglionic tract neuropiles. Golgi impregnation and intracellular dye filling show that local interneurons originate in ventral sensory neuropiles of leg neuromeres and ascend dorsally to terminate amongst dendrites of motor neurons. Local interneurons generally show higher thresholds for vibration stimuli than do receptors. Local interneurons typically receive inputs from one or several types of receptors. Some respond to stimulation of a single leg, others respond to stimulation of several legs on the same side of the body. The functional morphology of the receptor afferents is correlated with known physiological characteristics of slit sensilla and trichobothria. Structure and activity of the local interneurons are compared with analogous interneurons in other arthropods.     

The responses of central acoustic and vibratory interneurones in bushcrickets and locusts to ultrasonic stimulation

ABSTRACT. Analysis of the ultrasonic content of the calling songs of two tettigoniids, Decticus verrucivorus L. and Tettigonia cantans Fuessly, showed that the major secondary energy peaks in the ultrasonic range are only about 15–20 dB below the main audible frequency peaks. The song of the acridid, Locusta migratoria L., contains no appreciable secondary peaks at ultrasonic frequencies, Bifunctional acoustic-vibratory interneurones are present in the ventral nerve cord of all three species. They are divided into three categories, according to their response characteristics: VS (vibration and sound), S (sound) and V (vibration) neurones. All the unit-types capable of coding sound signals in the ventral cord (VS and S neurones) are sensitive to frequencies of up to 100 kHz, with one exception (S3). In tettigoniids, three of these unit-types are more sensitive at ultrasonic frequencies than they are at the audible frequencies of their conspecific songs. Among the vibratory neurones (V), one unit-type receives inhibitory inputs from ultrasonic acoustic primary receptors. The possible importance of ultrasonic perception in the natural environment is briefly discussed.     

Ein auf Substratvibration reagierendes Interneuron im Bauchmark der Grille

Summary By extracellular recording from the neck connectives of free moving crickets potentials of large fibres can be obtained, which respond to substratum vibration. The most sensitive fibres which seem to be connected to the subgenual organs show an adaptation which can be modified by central or peripheral factors. The sensitivity of one fibre which was tested with a vibration platform of adjustable frequency and amplitude has a threshold of 0.2 to 0.3 cm/sec² of acceleration at a frequency of 500 Hz (Fig. 4). These data correspond with the results of threshold measurements on the subgenual organs made by Autrum and Schneider (1948).

---

---

Die Untersuchungen wurden unterstützt durch Mittel der Deutschen Forschungsgemeinschaft, die Herrn Prof. Dr. F. Huber zur Verfügung standen.     

Spiders of the genus Cupiennius Simon 1891 (Araneae,Ctenidae)

Summary Cupiennius salei (Ctenidae) is a tropical wandering spider which lives in close association with a particular type of plant (see companion paper). These plants are the channels through which the spiders receive and emit various types of vibrations. We measured the vibrations the spiders are typically exposed to when they sit on their dwelling plants (banana plant, bromeliad) in their natural biotope in Central America. In addition a laboratory analysis was carried out to get an approximate idea of the complex vibration-propagating properties of the dwelling plants, taking a banana plant as an example. (1) Types of vibrations (Figs. 1–4). Despite variability in detail there are characteristic differences in spectral composition between the vibrations of various abiotic and biotic origins: (a) Vibrations due to wind are very low frequency phenomena. Their frequency spectra are conspicuously narrow with prominent peaks close to or, more often, below 10 Hz. Vibrations due to raindrops show maximal acceleration values at ca. 1000 Hz. Their frequency band at-20 dB extends up to ca. 250 Hz where-as that of the vibrations due to wind extends to only ca. 50 Hz. (b) The frequency spectra of prey vibrations such as those generated by a running cockroach are typically broad-banded and contain high frequencies; they have largest peaks mostly between ca. 400 and 900 Hz. Their-20 dB frequency bands usually extend from a few Hz to ca. 900 Hz. Some potential prey animals such as grass-hoppers seem to be vibrocryptic; they walk by the spider as if unnoticed. Their cautious gait leads to only weak vibrations at very low frequencies resembling the background noise due to wind. Courtship signals are composed maily of low frequencies, intermediate between background noise and prey vibrations (male: prominent peaks at ca. 75 Hz and ca. 115 Hz; female: dominant frequencies between ca. 20 Hz and ca. 50 Hz). The male signal is composed of syllables and differs from all other vibrations studied here by being temporally highly ordered. A comparison with previous electrophysiological studies suggests that the high pass characteristics of the vibration receptors enhance the signal-to-(abiotic)-noise ratio and that the vibration-sensitive interneurons so far examined and found to have band pass characteristics are tuned to the frequencies found in the vibrations of biotic origin. (2) Signal propagation (Fig. 5). In terms of frequency-dependent attenuation of vibrations the banana plant is well suited for transmitting the above signals. Average attenuation values are ca. 0.35 dB/cm. Together with known data on vibration receptor sensitivity this explains the range of courtship signals of more than 1 m observed in behavioral studies. Attenuation in the plant is neither a monotonic function of frequency nor of distance from the signal source.     

‘Peripheral inhibition’ of vibration-sensitive units in the leg of the fiddler crabUca pugilator

Summary The activity of single vibration-sensitive neurons in the leg nerve of the fiddler crabUca pugilator was recorded extracellularly. All units recorded from fall into two groups according to basic differences in their spectral threshold curves. The first type of neuron can be excited over a broad frequency range (ca. 2–2,000 Hz) with minimal threshold at 15–30 Hz with 0.5–1.0 cm/s² (peak). The second type of neuron, in contrast to the first one spontaneously active, is excited only in the frequency range 2–100 Hz and shows a decrease in the nerve impulse rate at vibration frequencies up to 2 kHz. The intensity necessary for complete suppression of the firing activity is 80 cm/s² at 800 Hz, the range of frequency most sensitive for inhibition.     

Sound and vibration sensitivity of VIIIth nerve fibers in the grassfrog, Rana temporaria

We have studied the sound and vibration sensitivity of 164 amphibian papilla fibers in the VIIIth nerve of the grassfrog, Rana temporaria. The VIIIth nerve was exposed using a dorsal approach. The frogs were placed in a natural sitting posture and stimulated by free-field sound. Furthermore, the animals were stimulated with dorso-ventral vibrations, and the sound-induced vertical vibrations in the setup could be canceled by emitting vibrations in antiphase from the vibration exciter. All low-frequency fibers responded to both sound and vibration with sound thresholds from 23 dB SPL and vibration thresholds from 0.02 cm/s². The sound and vibration sensitivity was compared for each fiber using the offset between the rate-level curves for sound and vibration stimulation as a measure of relative vibration sensitivity. When measured in this way relative vibration sensitivity decreases with frequency from 42 dB at 100 Hz to 25 dB at 400 Hz. Since sound thresholds decrease from 72 dB SPL at 100 Hz to 50 dB SPL at 400 Hz the decrease in relative vibration sensitivity reflects an increase in sound sensitivity with frequency, probably due to enhanced tympanic sensitivity at higher frequencies. In contrast, absolute vibration sensitivity is constant in most of the frequency range studied. Only small effects result from the cancellation of sound-induced vibrations. The reason for this probably is that the maximal induced vibrations in the present setup are 6–10 dB below the fibers' vibration threshold at the threshold for sound. However, these results are only valid for the present physical configuration of the setup and the high vibration-sensitivities of the fibers warrant caution whenever the auditory fibers are stimulated with free-field sound. Thus, the experiments suggest that the low-frequency sound sensitivity is not caused by sound-induced vertical vibrations. Instead, the low-frequency sound sensitivity is either tympanic or mediated through bone conduction or sound-induced pulsations of the lungs.Abbreviations AP amphibian papilla - BF best frequency - PST peristimulus time     

Vibration receptive sensilla on the wing margins of the silkworm moth Bombyx mori

Bristles along the wing margins (wm-bristles) of the silkworm moth, Bombyx mori, were studied morphologically and electrophysiologically. The male moth has ca. 50 wm-bristles on each forewing and hindwing. Scanning electron microscopy revealed that these wm-bristles are typical mechanosensilla. Leuco-methylene blue staining demonstrated that each wm-bristle has a single receptor neuron, which is also characteristic of the mechanosensillum. The receptor neuron responded to vibrating air currents but did not respond to a constant air current. The wm-bristles showed clear directional sensitivity to vibrating air currents. The wm-bristles were classified into two types, type I and type II, by their response patterns to sinusoidal movements of the bristle. The neuron in type I discharged bursting spikes immediately following stimulation onset and also discharged a single spike for each sinusoidal cycle for frequencies less than ca. 60 Hz. The neuron in type II only responded to vibrations over 40 Hz and, specifically at 75 Hz, discharged a single spike for each sinusoidal cycle throughout the stimulation period. These results suggest that the two types of wm-bristles are highly tuned in different ways to detect vibrations due to the wing beat. The roles of the wm-bristles in the wing beat are discussed.     

Effects of mechanical vibration on seed germination of Arabidopsis thaliana (L.) Heynh

The effects of sinusoidal vibration (40-120 Hz, amplitude equal to or smaller than 0.42 mm) on seed germination of Arabidopsis thaliana were examined. When the amplitude of vibration was fixed at 0.42 mm, vibration with frequencies higher than 70 Hz increased the rate of seed germination. When the frequency of vibration was fixed at 100 Hz, vibration with amplitudes larger than 0.33 mm also increased the rate of germination. The increase in the rate of germination appeared dependent on acceleration calculated from the frequency and amplitude of vibration. Vibration with a maximum acceleration of 70 m s(-2) increased the rate of germination, but the promotive effects leveled off at higher accelerations. Vibration had little effect on seed germination in a starch-deficient mutant, pgm. Thus, the amyloplasts appeared to act as a susceptor that senses mechanical vibrations. No vibration-induced promotion of germination was seen in an ethylene-insensitive mutant, etr1, or in the wild type in the presence of aminoethoxyvinylglycine, an inhibitor of ethylene synthesis, suggesting that vibration increased the rate of seed germination through the action of ethylene.     

Modelling the subgenual organ of the honeybee, Apis mellifera

In a recent study on the honeybee (Apis mellifera), the subgenual organ was observed moving inside the leg during sinusoidal vibrations of the leg (Kilpinen and Storm 1997). The subgenual organ of the honeybee is suspended in a haemolymph channel in the tibia of each leg. When the leg accelerates, the inertia causes the haemolymph and the subgenual organ to lag behind the movement of the rest of the leg. To elucidate the biophysics of the subgenual organ system of the honeybee, two mathematical models to simulate the experimentally observed mechanical response are considered. The models are a classical mass-spring model and a newly developed tube model consisting of an open-ended, fluid-filled tube occluded by an elastic structure midway. Both models suggest that the subgenual organ included in the haemolymph channel resembles that of an overdamped system. In resembling the biophysics of the subgenual organ system in the honeybee, we consider the tube model to be the better of the two because it simulates a mechanical response which complies best with the experimental data, and the physical parameters in the model can be related to the␣constituent parts of the subgenual organ included in the haemolymph channel. Received: 25 July 1997 / Accepted in revised form: 8 December 1997     

The auditory system of blood-sucking mosquito females (Diptera, Culicidae): Acoustic perception during flight simulation

Mosquitoes hear with their plumose antennae which respond to the air movement caused by sound propagation and conduct vibrations to the Johnston’s organ located at the base of each antenna. Each of the two Johnston’s organs contains several tens of thousands mechanosensory cells which detect the displacements of the flagellum and transform them into electric potentials. Hearing plays a very important role in the reproductive behavior of the male mosquitoes. At the same time, our knowledge of hearing in female mosquitoes is very limited and its functional significance is obscure. In this study we measured the auditory sensitivity of female mosquitoes and investigated how the flight conditions affect their hearing. We studied mosquitoes of three species: Anopheles messeae, Aedes excrucians, and Culex pipiens pipiens. The neuronal responses were recorded with a glass microelectrode from the antennal nerve and the deutocerebral interneurons. Stimulation was applied in two modes: (1) the main stimulus against the background of flight simulation (strong vibration with the typical wingbeat frequency of a given mosquito species) and (2) only the main stimulus without the background stimulation. During the flight simulation, females demonstrated an increased sensitivity to frequencies below 200 Hz. The mean auditory receptor threshold at 80–120 Hz was 45 dB, which was 8 dB lower than that without flight simulation. An additional zone of increased sensitivity was also found at frequencies higher than the simulated wingbeat frequency (the so-called image channel). Our analysis of frequency tuning curves measured from the receptors and auditory interneurons shows that mosquito auditory neuronal complex consists of several subsystems which have different frequency tuning parameters, and suggests the possibility of spectral analysis of sounds. Three hypotheses could be proposed on the function of hearing in female mosquitoes: (1) predator avoidance, (2) detection of moving prey, and (3) intraspecific communication. Each of the hypotheses involves the ability to analyze the sound frequency spectrum and subsequent signal recognition.     

A Functionally Equivalent Artificial Neural Network Model of the Prey Orientation Behavior of Waterstriders (Gerridae)

Waterstriders, a family (Heteroptera, Gerridae) of predacious insects, orient toward the source of water surface vibrations. We describe an artificial neural network that simulates a waterstrider's discrete rotational movement towards a prey item and compare the results to published data. A back-propagation network with six input units, each corresponding to a vibration receptor on a leg of the waterstrider, and two output units corresponding to the elicited angle of rotation, was used. The network was trained with a full complement of receptors to rotate towards the point source of a surface vibration. When the network was tested with all receptors present, a linear relationship was found between the desired and obtained rotational angles. Lesioning of one or two receptors resulted in marked deviation from linearity within the angular range of detection corresponding to that of the amputated receptor(s), while amputation of three receptors resulted in the network rotating contralaterally to all vibrations originating ipsolaterally to the lesioned side. All trials produced results that corresponded qualitatively to published behavioral data.