The antennal ear of the fruit fly, called the Johnston's organ (JO), detects a wide variety of mechanosensory stimuli, including sound, wind, and gravity. Like many sensory cells in insect, JO ...neurons are compartmentalized in a sensory unit (i.e., scolopidium). To understand how different subgroups of JO neurons are organized in each scolopidial compartment, we visualized individual JO neurons by labeling various subgroups of JO neurons in different combinations. We found that vibration-sensitive (or deflection-sensitive) neurons rarely grouped together in a single scolopidial compartment. This finding suggests that JO neurons are grouped in stereotypical combinations each with a distinct response property in a scolopidium.
Projections from peripheral receptors directly into the protocerebrum of insects have only been little studied. Retrograde staining of nerves from the antennae, maxillary palps and legs has revealed ...some fibres that project into the central areas of the protocerebrum. In the case of the antennae and palps, it was not known which receptors were responsible for these projections. In the legs of locusts, multipolar neurons (MN) with characteristic terminal dendritic masses (TDM) have been described to project into a neuropil called “superior ventral inferior protocerebrum” (SVIP). However, such neurons have only been found in the abdominal infrared organs of the Australian fire beetle
Merimna atrata
, where they function as thermoreceptors. In several orthopterans, fibres from the antennae and palps also project into the SVIP. The present work suggests that the multipolar neuron from the infrared organ of
Merimna
also projects into the protocerebrum, possibly into a ventral region functionally analogous to the SVIP. No MNs but single scolopidia were found in the tips of the antennae and palps of locusts, apparently responsible for projections into the SVIP, where they probably function as receptors for haemolymph pressure.
Centipedes are terrestrial, predatory arthropods with specialized sensory organs. However, many aspects of their sensory biology are still unknown. This also concerns hygroreception, which is ...especially important for centipedes, as their epicuticle is thin and they lose water rapidly at low humidity. Thus, the detection of humid places is vital but to date no definite hygroreceptor was found in centipedes. House centipedes (Scutigeromorpha) possess a peculiar opening at the base of their antenna, termed 'scape organ', that houses up to 15 cone-shaped sensilla in a cavity. Lacking wall and tip-pores, these socket-less sensilla may be hypothesized to function as hygroreceptors similar to those found in hexapods.
The cone-shaped sensilla in the scape organ as well as nearby peg-shaped sensilla are composed of three biciliated receptor cells and three sheath cells. A tip-pore is present but plugged by a highly electron-dense secretion, which also overlays the entire inner surface of the cavity. Several solitary recto-canal epidermal glands produce the secretion. Receptor cell type 1 (two cells in cone-shaped sensilla, one cell in peg-shaped sensilla) possesses two long dendritic outer segments that project to the terminal pore. Receptor cell type 2 (one cell in both sensilla) possesses two shorter dendritic outer segments connected to the first (proximal) sheath cell that establishes a scolopale-like structure, documented for the first time in detail in a myriapod sensillum.
The nearly identical configuration of receptor cells 1 with their long dendritic outer segments in both sensilla is similar to hexapod hygroreceptors. In Scutigera coleoptrata, however, the mechanism of stimulus transduction is different. Water vapor may lead to swelling and subsequent elongation of the plug pin that enters the terminal pore, thus causing stimulation of the elongated dendritic outer segments. The interconnection of receptor cell 2 with short outer dendritic segments to a scolopale-like structure potentially suits both sensilla for vibration or strain detection. Thus, both sensilla located at the antennal base of scutigeromorph centipedes fulfill a dual function.
Ion homeostasis is a fundamental cellular process particularly important in excitable cell activities such as hearing. It relies on the Na ⁺/K ⁺ ATPase (also referred to as the Na pump), which is ...composed of a catalytic α subunit and a β subunit required for its transport to the plasma membrane and for regulating its activity. We show that α and β subunits are expressed in Johnston's organ (JO), the Drosophila auditory organ. We knocked down expression of α subunits (ATPα and α-like) and β subunits (nrv1, nrv2 , and nrv3) individually in JO with UAS/Gal4-mediated RNAi. ATPα shows elevated expression in the ablumenal membrane of scolopale cells, which enwrap JO neuronal dendrites in endolymph-like compartments. Knocking down ATPα , but not α-like , in the entire JO or only in scolopale cells using specific drivers, resulted in complete deafness. Among β subunits, nrv2 is expressed in scolopale cells and nrv3 in JO neurons. Knocking down nrv2 in scolopale cells blocked Nrv2 expression, reduced ATPα expression in the scolopale cells, and caused almost complete deafness. Furthermore, knockdown of either nrv2 or ATPα specifically in scolopale cells causes abnormal, electron-dense material accumulation in the scolopale space. Similarly, nrv3 functions in JO but not in scolopale cells, suggesting neuron specificity that parallels nrv2 scolopale cell–specific support of the catalytic ATPα. Our studies provide an amenable model to investigate generation of endolymph-like extracellular compartments.
Insect ears evolved many times independently. As a consequence, a striking diversity exists in the location, construction and behavioural implementation of ears. In this review, we first summarise ...what is known about the evolutionary origin of ears and the presumed precursor organs in the various insect groups. Thereafter, we focus on selective forces for making and keeping an ear: we discuss detecting and localising predators and conspecifics, including establishing new “private” channels for intraspecific communication. More advanced aspects involve judging the distance of conspecifics, or assessing individual quality from songs which makes auditory processing a means for exerting sexual selection on mating partners. We try to identify negative selective forces, mainly in the context of energy expenditure for developing and keeping an ear, but also in conjunction with acoustic communication, which incorporates risks like eavesdropping by predators and parasitoids. We then discuss balancing pressures, which might oppose optimising an ear for a specific task (when it serves different functions, for example). Subsequently, we describe various scenarios that might have led to a reduction or complete loss of ears in evolution. Finally, we describe cases of sex differences in ears and potential reasons for their appearance.
The tympanal organ of the bushcricket Mecopoda elongata emits pronounced distortion-product otoacoustic emissions (DPOAEs). Their characteristics are comparable to those measured in other insects, ...such as locusts and moths, with the 2f1-f2 emission being the most prominent one. Yet the site of their generation is still unclear. The spatial separation between the sound receiving spiracle and the hearing organ in this species allows manipulations of the sensory cells without interfering with the acoustical measurements. We tried to interfere with the DPOAE generation by pharmacologically influencing the tympanal organ using the insecticide pymetrozine. The compound appears to act selectively on scolopidia, i.e., the mechanosensor type characteristically constituting tympanal organs. Pymetrozine solutions were applied as closely as possible to the scolopidia via a cuticle opening in the tibia, distally to the organ. Applications of pymetrozine at concentrations between 10⁻³ and 10⁻⁷ M to the tympanal organ led to a pronounced and irreversible decrease of the DPOAE amplitudes.
Most individuals of the Australian ‘fire-beetle’
Merimna atrata have two pairs of IR receptors which are located ventrolaterally on the second and third abdominal sternite. An IR receptor consists of ...a specialized IR absorbing area, which is innervated by a neural complex. This complex contains one thermoreceptive multipolar neuron with a unique terminal dendritic mass (TDM) and two scolopidia and was termed ‘sensory complex’. However, also individuals with one pair of IR receptors on the second sternite and beetles with three pairs on the second, third, and fourth sternites were found. Additionally, beetles having one or two pairs of IR receptors may have preliminary stages of IR receptors on the third and fourth sternite, respectively. We found two kinds of preliminary stages, both of which are characterized by a much less pronounced absorbing area. In all five abdominal sternites segmental nerves are attached to the cuticle with a neural complex. Investigation of complexes of non-IR sternites suggests that the sensory cells inside the sensory complex of an IR receptor have developed from common internal stretch receptors. From our results it can be hypothesized that the IR sensory system in
Merimna atrata has not yet reached a stage, which can be regarded as evolutionary stable.
The femoral chordotonal organ (FCO) inChrysoperla carneais situated in the distal part of the femur and consists of two scoloparia, which are fused at their distal end. The distal scoloparium ...contains 17–20 scolopidia, and the proximal one six scolopidia. Each scolopidium consists of two sensory cells and three types of enveloping cells (glial, scolopale and attachment cell). The sensory cells of different scolopidia do not lie at the same level in the FCO. Therefore the attachment cells of different scolopidia have different lengths. In the FCO, three types of ciliary roots are found in different sensory cells. The dendrite of the sensory cell terminates in a distal process, which has the structure of a modified cilium (9×2+0). The very distal part of the cilium is surrounded by an extracellular electron dense material, the cap, and ends in a terminal dilation. The scolopale cell contains the electron dense scolopale rods, consisting of plentiful microtubules. In their middle third the scolopale rods are fused and form the scolopale. In the FCO septate junctions, desmosomes and hemidesmosomes are found.
More than 50 chordotonal sensilla, or scolopidia, embedded entirely in the integument were found in each side of the genital chamber wall in the female cricket, Teleogryllus commodus. Their cell ...bodies lie among the epidermal cells, and the tips of their dendrites terminate in the cuticle. About half of them contain two sensory cells (two-cell scolopidium), the others only one (one-cell scolopidium). The sensory cell in the one-cell scolopidium is the type-1 cell. In the two-cell scolopidium one is type-1 and the other type-2. Regardless of the number of sensory cells, they are all amphinematic. In the two-cell scolopidium only the type-2 dendrite, rich in microtubules, penetrates into the cuticle, bifurcates and terminates in the tube enclosed by an attachment cell; the type-1 never extends into the cuticle. On the other hand, the type-1 cell in the one-cell scolopidium projects its apex into the cuticle. The unique topography and structure of these scolopidia lead to the following hypothesis about the phylogenetic relationship between the scolopidia and other kinds of sensilla: the type-1 scolopidial sensory cell buried in the integument may be the original model, which through the loss of the long regular axoneme has given rise to type-2 cells. Modification of the apical region, the tubular body or ramification, may have lead to the cuticular sensilla corresponding to the development of the cuticular apparatus, and the scolopidia may have been withdrawn into the body cavity to form ordinary chordotonal organs.
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