Sensing of Substrate Vibrations in the Adult Cicada Okanagana rimosa (Hemiptera: Cicadidae)

Detection of substrate vibrations is an evolutionarily old sensory modality and is important for predator detection as well as for intraspecific communication. In insects, substrate vibrations are detected mainly by scolopidial (chordotonal) sense organs found at different sites in the legs. Among these sense organs, the tibial subgenual organ (SGO) is one of the most sensitive sensors. The neuroanatomy and physiology of vibratory sense organs of cicadas is not well known. Here, we investigated the leg nerve by neuronal tracing and summed nerve recordings. Tracing with Neurobiotin revealed that the cicada Okanagana rimosa (Say) (Hemiptera: Cicadidae) has a femoral chordotonal organ with about 20 sensory cells and a tibial SGO with two sensory cells. Recordings from the leg nerve show that the vibrational response is broadly tuned with a threshold of about 1 m/s2 and a minimum latency of about 6 ms. The vibratory sense of cicadas might be used in predator avoidance and intraspecific communication, although no tuning to the peak frequency of the calling song (9 kHz) could be found

Developmental constraint of insect audition

BACKGROUND: Insect ears contain very different numbers of sensory cells, from only one sensory cell in some moths to thousands of sensory cells, e.g. in cicadas. These differences still await functional explanation and especially the large numbers in cicadas remain puzzling. Insects of the different orders have distinct developmental sequences for the generation of auditory organs. These sensory cells might have different functions depending on the developmental stages. Here we propose that constraints arising during development are also important for the design of insect ears and might influence cell numbers of the adults. PRESENTATION OF THE HYPOTHESIS: We propose that the functional requirements of the subadult stages determine the adult complement of sensory units in the auditory system of cicadas. The hypothetical larval sensory organ should function as a vibration receiver, representing a functional caenogenesis. TESTING THE HYPOTHESIS: Experiments at different levels have to be designed to test the hypothesis. Firstly, the neuroanatomy of the larval sense organ should be analyzed to detail. Secondly, the function should be unraveled neurophysiologically and behaviorally. Thirdly, the persistence of the sensory cells and the rebuilding of the sensory organ to the adult should be investigated. IMPLICATIONS OF THE HYPOTHESIS: Usually, the evolution of insect ears is viewed with respect to physiological and neuronal mechanisms of sound perception. This view should be extended to the development of sense organs. Functional requirements during postembryonic development may act as constraints for the evolution of adult organs, as exemplified with the auditory system of cicadas

Embryonic development of pleuropodia of the cicada, Magicicada cassini

In many insects the first abdominal segment possesses embryonic appendages called pleuropodia. Here we show the embryogenesis of pleuropodial cells of the periodical cicada, Magicicada cassini (Fisher 1851) (Insecta, Homoptera, Cicadidae). An antibody, anti-horseradish perioxidase (HRP), that is usually neuron-specific strongly marked the pleuropodial anlagen and revealed their ectodermal origin shortly after limb bud formation. Thereafter the cells sank into the epidermis and their apical parts enlarged. A globular part protruded from the body wall. Filamentous structures were marked at the stem region and into the apical dilation. In later embryonic stages the pleuropodia degenerated. Despite the binding of anti-HRP the cells had no morphological neuronal characters and cannot be regarded as neurons. The binding indicates that glycosylated cell surface molecules contribute to the adhesion between the presumably glandular pleuropodial cells. In comparison, anti-HRP does not mark the pleuropodia of Orthoptera

Sounds, Behaviour, and Auditory Receptors of the Armoured Ground Cricket, Acanthoplus longipes

The auditory sensory system of the taxon Hetrodinae has not been studied previously. Males of the African armoured ground cricket, Acanthoplus longipes (Orthoptera: Tettigoniidae: Hetrodinae) produce a calling song that lasts for minutes and consists of verses with two pulses. About three impulses are in the first pulse and about five impulses are in the second pulse. In contrast, the disturbance stridulation consists of verses with about 14 impulses that are not separated in pulses. Furthermore, the inter-impulse intervals of both types of sounds are different, whereas verses have similar durations. This indicates that the neuronal networks for sound generation are not identical. The frequency spectrum peaks at about 15 kHz in both types of sounds, whereas the hearing threshold has the greatest sensitivity between 4 and 10 kHz. The auditory afferents project into the prothoracic ganglion. The foreleg contains about 27 sensory neurons in the crista acustica; the midleg has 18 sensory neurons, and the hindleg has 14. The auditory system is similar to those of other Tettigoniidae

Adaptive Strategies in Life-History of Bushcrickets (Orthoptera) and Cicadas (Homoptera) to Parasitoids Pressure on Their Acoustic Communication Systems—A Case for Sociality?

In sexual reproduction, the search for mating partners elevates the individual’s risks of predation and parasitism. One way to increase mate search effectiveness and reduce search costs is acoustic signaling. However, acoustic orienting parasitoid flies exploit singing hosts, leading to high parasitism rates. Aggregations of males and females at mating and singing in choruses might reduce individual risks by dilution and predator saturation. This mini-review reflects on consequences for host’s acoustic signaling in choruses using the examples of cicadas and bushcrickets. It concludes that despite antagonistic selection pressure by parasitoids, singing in choruses might select for increased, not reduced signaling in males. The time joining and leaving a chorus might be crucial: once mated, a refractory period will drop males off the signaling pool, preventing parasitism. In a chorus, fast and loud singing might be highly advantageous, supporting the fittest males. Natural selection might have shaped signaling strategies in choruses, which can probably only be understood when applying individual based dynamic modeling.Peer Reviewe

Contralateral Deafferentation Does Not Affect Regeneration Processes in the Auditory System of (Orthoptera)

The auditory system of locusts has high regeneration capacity following injury of the peripheral afferents. Regenerating auditory afferents can re-innervate their target areas even after changed neuronal pathways. Here, possible influences of contralateral deafferentation on regenerating afferents were investigated. Contralateral deafferentation was performed at different stages of the regeneration. Regeneration was triggered by crushing the tympanal nerve. The regenerated fibers showed aberrant fiber outgrowth, reduced density of terminations in the target area, the auditory neuropile and collateral sprouts crossing the midline. However, these results were not significantly influenced by the contralateral deafferentation. Therefore the bilateral symmetrical systems seem to be largely independent from each other

The subgenual organ complex in the cave cricket Troglophilus neglectus (Orthoptera: Rhaphidophoridae)

Comparative studies of the organization of nervous systems and sensory organs can reveal their evolution and specific adaptations. In the forelegs of some Ensifera (including crickets and tettigoniids), tympanal hearing organs are located in close proximity to the mechanosensitive subgenual organ (SGO). In the present study, the SGO complex in the non-hearing cave cricket Troglophilus neglectus (Rhaphidophoridae) is investigated for the neuronal innervation pattern and for organs homologous to the hearing organs in related taxa. We analyse the innervation pattern of the sensory organs (SGO and intermediate organ (IO)) and its variability between individuals. In T. neglectus, the IO consists of two major groups of closely associated sensilla with different positions. While the distal-most sensilla superficially resemble tettigoniid auditory sensilla in location and orientation, the sensory innervation does not show these two groups to be distinct organs. Though variability in the number of sensory nerve branches occurs, usually either organ is supplied by a single nerve branch. Hence, no sensory elements clearly homologous to the auditory organ are evident. In contrast to other non-hearing Ensifera, the cave cricket sensory structures are relatively simple, consistent with a plesiomorphic organization resembling sensory innervation in grasshoppers and stick insects

Data from: Position-dependent hearing in three species of bushcrickets (Tettigoniidae, Orthoptera)

A primary task of auditory systems is the localization of sound sources in space. Sound source localization in azimuth is usually based on temporal or intensity differences of sounds between the bilaterally arranged ears. In mammals, localization in elevation is possible by transfer functions at the ear, especially the pinnae. Although insects are able to locate sound sources, little attention is given to the mechanisms of acoustic orientation to elevated positions. Here we comparatively analyse the peripheral hearing thresholds of three species of bushcrickets in respect to sound source positions in space. The hearing thresholds across frequencies depend on the location of a sound source in the three-dimensional hearing space in front of the animal. Thresholds differ for different azimuthal positions and for different positions in elevation. This position-dependent frequency tuning is species specific. Largest differences in thresholds between positions are found in Ancylecha fenestrata. Correspondingly, A. fenestrata has a rather complex ear morphology including cuticular folds covering the anterior tympanal membrane. The position-dependent tuning might contribute to sound source localization in the habitats. Acoustic orientation might be a selective factor for the evolution of morphological structures at the bushcricket ear and, speculatively, even for frequency fractioning in the ear