Functional Morphology of Leg Mechanosensory Organs in Early Postembryonic Development in the Stick Insect (Sipyloidea chlorotica).

The subgenual organ complex of stick insects has a unique neuroanatomical organisation with two elaborate chordotonal organs, the subgenual organ and the distal organ. These organs are present in all leg pairs and are already developed in newly hatched stick insects. The present study analyses for the first time the morphology of sensory organs in the subgenual organ complex for a membrane connecting the two sensory organs in newly hatched insects (Sipyloidea chlorotica (Audinet-Serville 1838)). The stick insect legs were analysed following hatching by axonal tracing and light microscopy. The subgenual organ complex in first juvenile instars shows the sensory organs and a thin membrane connecting the sensory organs resembling the morphology of adult animals. Rarely was this membrane not detected, where it is assumed as not developed during embryogenesis. The connection appears to influence the shape of the subgenual organ, with one end extending towards the distal organ as under tension. These findings are discussed for the following functional implications: (1) the physiological responses of the subgenual organ complex to mechanical stimuli after hatching, (2) the influence of the membrane on the displacement of the sensory organs, and (3) the connection between the subgenual organ and distal organ as a possible functional coupling.

Vibration receptor organs in the insect leg: neuroanatomical diversity and functional principles.

Detecting substrate vibrations is essential for insects in different behavioural contexts. These vibrational behaviours are mediated by mechanoreceptor organs detecting and processing vibrational stimuli transmitted in the environment. We discuss recently gained insights about the functional principles of insect vibration receptors, mainly leg chordotonal organs highly sensitive to vibrational stimuli, and the mechanisms of their diversification in neuroanatomy and functional morphology, in relation to the attachment structures and mechanical coupling. The two main input pathways for vibration stimuli transferred by the insect legs to vibrosensory organs via the cuticle and via the hemolymph are fundamental for explaining sensory specialisations. The vibroreceptor organs can diversify in their neuroanatomy and morphology in several key aspects. This provides the structural basis for complex adaptations in sensory evolution.

The neuronal innervation pattern of the subgenual organ complex in Peruphasma schultei (Insecta: Phasmatodea).

The proximal tibia of orthopteroid insects contains sensory organs, the subgenual organ complex, detecting mechanical stimuli including substrate vibration. In stick insects, two chordotonal organs occur in close proximity, the subgenual organ and the distal organ, which likely detect substrate vibrations. In most stick insects, both organs are innervated by separate nerve branches. To obtain more data on the neuroanatomy of the subgenual organ complex from the New World phasmids (Occidophasmata), the present study documents the neuronal innervation of sensory organs in the subgenual organ complex of Peruphasma schultei, the first species from Pseudophasmatinae investigated for this sensory complex. The innervation pattern shows a distinct nerve branch for the subgenual organ and for the distal organ in most cases. Some variability in the innervation, which generally occurs for these chordotonal organs, was noted for both organs in P. schultei. The most common innervation for both organs was by a single nerve branch for each organ. The innervation of the subgenual organ resembled the nerve pattern of another New World phasmid, but was simpler than in the Old World phasmids (Oriophasmata) studied so far. Therefore, the peripheral neuronal innervation of sensory organs could reflect phylogenetic relationships and provide phylogenetic information, while the overall neuroanatomy of the subgenual organ complex is similar in stick insects.

Comparative Neuroanatomy of the Mechanosensory Subgenual Organ Complex in the Peruvian Stick Insect, Oreophoetes peruana.

The subgenual organ complex in the leg of Polyneoptera (Insecta) consists of several chordotonal organs specialized to detect mechanical stimuli from substrate vibrations and airborne sound. In stick insects (Phasmatodea), the subgenual organ complex contains the subgenual organ and the distal organ located distally to the subgenual organ. The subgenual organ is a highly sensitive detector for substrate vibrations. The distal organ has a characteristic linear organization of sensilla and likely also responds to substrate vibrations. Despite its unique combination of sensory organs, the neuroanatomy of the subgenual organ complex of stick insects has been investigated for only very few species so far. Phylogenomic analysis has established for Phasmatodea the early branching of the sister groups Oriophasmata, the Old World phasmids, and Occidophasmata, the New World phasmids. The species studied for the sensory neuroanatomy, including the Indian stick insect Carausius morosus, belong to the Old World stick insects. Here, the neuroanatomy of the subgenual organ complex is presented for a first species of the New World stick insects, the Peruvian stick insect Oreophoetes peruana. To document the sensory organs in the subgenual organ complex and their innervation pattern, and to compare these between females and males of this species and also to the Old World stick insects, axonal tracing is used. This study documents the same sensory organs for O. peruana, subgenual organ and distal organ, as in other stick insects. Between the sexes of this species, there are no notable differences in the neuroanatomy of their sensory organs. The innervation pattern of tibial nerve branches in O. peruana is identical to other stick insect species, although the innervation pattern of the subgenual organ by a single tibial nerve branch is simpler. The shared organization of the organs in the subgenual organ complex in both groups of Neophasmatodea (Old World and New World stick insects) indicates the sensory importance of the subgenual organ but also of the distal organ. Some variation exists in the innervation of the chordotonal organs in O. peruana though a common innervation pattern can be identified. The findings raise the question for the ancestral neuroanatomical organization and innervation in stick insects.

Vibration detection in arthropods: Signal transfer, biomechanics and sensory adaptations.

In arthropods, the detection of vibrational signals and stimuli is essential in several behaviours, including mate recognition and pair formation, prey detection, and predator evasion. These behaviours have been studied in several species of insects, arachnids, and crustaceans for vibration production and propagation in the environment. Vibration stimuli are transferred over the animals' appendages and the body to vibrosensory organs. Ultimately, the stimuli are transferred to act on the dendrites of the mechanosensitive sensilla. We refer to these two different levels of transfer as macromechanics and micromechanics, respectively. These biomechanical processes have important roles in filtering and pre-processing of stimuli, which are not carried out by neuronal components of sensory organs. Also, the macromechanical transfer is posture-dependent and enables behavioural control of vibration detection. Diverse sensory organs respond to vibrations, including cuticular sensilla (slit sensilla, campaniform sensilla) and internal chordotonal organs. These organs provide various adaptations, as they occur at diverse body positions with different mechanical couplings as input pathways. Macromechanics likely facilitated evolution of vibrosensory organs at specific body locations. Thus, vibration detection is a highly complex sensory capacity, which employs body and sensory mechanics for signal filtering, amplification, and analysis of frequency, intensity and directionality.

Are all auditory sensilla of bushcrickets bimodal? Comment on: R. D. Zhantiev and O. S. Korsunovskaya, Functions of chordotonal sensilla in bushcrickets (Orthoptera, Tettigoniidae); Entomological Review, 2021, vol. 101 (6), pp. 755-766.

Detection of sound and substrate vibration is crucial for the survival and reproduction of many animals, particularly insects. Bushcrickets (Orthoptera, Tettigoniidae), developed a large mechanosensory organ complex in their legs to detect such stimuli. As demonstrated by various studies in the past, sensilla in distinct functional groups form specialized vibratory organs (the subgenual organ and the accessory organ), respond sensitively to both vibration and sound (in the intermediate organ [IO]), or mediate hearing (in the crista acustica [CA]; the tympanal hearing organ). In their recent publication, Zhantiev and Korsunovskaya addressed auditory and vibratory sensitivity in the IO and the CA in two species of bushcrickets, using single-cell recording and staining of sensory neurons from their soma in an isolated foreleg. Their main finding was that not only the IO but also the complete CA contains bimodal sensilla responding with high sensitivity to both sound and vibration, which would be a true change in the paradigm of how the auditory/vibratory sense in Orthoptera works. In addition, they revealed vibratory tuning of the IO sensilla, which differs largely from that in previous studies. We propose three major experimental causes of such discrepancies: calibration, experiments with isolated legs, and differences in the sites of recording. To judge the causes of these discrepancies more adequately, a detailed comparison of methods and a number of control experiments are needed. This will deepen our understanding of sensory adaptations and specialization of insect mechanosensory organs to stimuli entering the system by different input pathways.

The tracheal system in the stick insect prothorax and prothoracic legs: Homologies to Orthoptera and relations to mechanosensory functions.

Arthropod respiration depends on the tracheal system running from spiracles at the body surface through the body and appendages. Here, three species of stick insects (Carausius morosus, Ramulus artemis, Sipyloidea sipylus) are investigated for the tracheae in the prothorax and foreleg. The origin of the tracheae from the mesothoracic spiracle that enter the foreleg is identified: five tracheae originate from the mesothoracic spiracle, of which two enter the foreleg (supraventral trachea, trachea pedalis anterior). These two tracheae run separately through the leg to the femur-tibia joint where they fuse, but in the proximal tibia split again into two tracheae. The leg tracheae in stick insects are homologous to those in Tettigoniidae (bushcrickets). Stick insects have two chordotonal organs in the proximal tibia (subgenual organ and distal organ) which locate dorsally of the leg trachea. The tracheal system shows no adaptation specific to the propagation of airborne sound, like enlarged spiracles or tracheal volumes. Tracheal vesicles form in the tibia proximally to the mechanosensory organs, but no tracheal sacks or expansions occur at the level of the sensory organs that could mediate the detection of airborne sound or amplify substrate vibrations transmitted in the hemolymph fluid. Rather, the morphological characteristics indicate a respiratory function.

Early postembryogenic development of the subgenual organ complex in the stick insect Sipyloidea sipylus.

Stick insects have elaborate mechanosensory organs in their subgenual organ complex in the proximal tibia, particularly the distal organ with scolopidial sensilla in linear arrangement. For early postembryonic developmental stages of Sipyloidea sipylus (Phasmatodea: Necrosciinae), the neuroanatomy of the scolopidial organs in the subgenual organ complex and the campaniform sensilla is documented by retrograde axonal tracing, and compared to the adult neuroanatomy. Already after hatching of the first larval instars are the sensory structures of subgenual organ and distal organ as well as tibial campaniform sensilla differentiated. In the distal organ, the full set of sensilla is shown in all larval stages examined. This finding indicates that the sensory organs differentiate during embryogenesis, and are already functional by the time of hatching. The constancy of distal organ sensilla over postembryonic stages allows investigation of the representative number of sensilla in adult animals as well as in larval instars. Some anatomical changes occur by postembryogenic length increase of the distal organ, and grouping of the anterior subgenual sensilla. The embryonic development of scolopidial sensilla is similar for auditory sensilla in hemimetabolous Orthoptera (locusts, bushcrickets, crickets) where tympanal membranes develop during postembryogenic stages, conferring a successive gain of sensitivity with larval moults.

Neuronal Innervation of the Subgenual Organ Complex and the Tibial Campaniform Sensilla in the Stick Insect Midleg.

Mechanosensory organs in legs play are crucial receptors in the feedback control of walking and in the detection of substrate-borne vibrations. Stick insects serve as a model for the physiological role of chordotonal organs and campaniform sensilla. This study documents, by axonal tracing, the neural innervation of the complex chordotonal organs and groups of campaniform sensilla in the proximal tibia of the midleg in Sipyloidea sipylus. In total, 6 nerve branches innervate the different sensory structures, and the innervation pattern associates different sensilla types by their position. Sensilla on the anterior and posterior tibia are innervated from distinct nerve branches. In addition, the variation in innervation is studied for five anatomical branching points. The most common variation is the innervation of the subgenual organ sensilla by two nerve branches rather than a single one. The fusion of commonly separated nerve branches also occurred. However, a common innervation pattern can be demonstrated, which is found in >75% of preparations. The variation did not include crossings of nerves between the anterior and posterior side of the leg. The study corrects the innervation of the posterior subgenual organ reported previously. The sensory neuroanatomy and innervation pattern can guide further physiological studies of mechanoreceptor organs and allow evolutionary comparisons to related insect groups.

Beyond the display: phase-only liquid crystal on Silicon devices and their applications in photonics [Invited].

Existing for almost four decades, liquid crystal on Silicon (LCOS) technology is rapidly growing into photonic applications. We review the basics of the technology, from the wafer to the driving solutions, the progress over the last decade and the future outlook. Furthermore we review the most exciting industrial and scientific applications of the LCOS technology.

Low-frequency vibration transmission and mechanosensory detection in the legs of cave crickets.

Vibrational communication is common in insects and often includes signals with prominent frequency components below 200 Hz, but the sensory adaptations for their detection are scarcely investigated. We performed an integrative study of the subgenual organ complex in Troglophilus cave crickets (Orthoptera: Rhaphidophoridae), a mechanosensory system of three scolopidial organs in the proximal tibia, for mechanical, anatomical and physiological aspects revealing matches to low frequency vibration detection. Microcomputed tomography shows that a part of the subgenual organ sensilla and especially the accessory organ posteriorly in this complex are placed closely underneath the cuticle, a position suited to evoke responses to low-frequency vibration via changes in the cuticular strain. Laser-Doppler vibrometry shows that in a narrow low-frequency range the posterior tibial surface reacts stronger to low frequency sinusoidal vibrations than the anterior tibial surface. This finding suggests that the posterior location of sensilla in tight connection to the cuticle, especially in the accessory organ, is adapted to improve detectability of low-frequency vibration signals. By electrophysiological recordings we identify a scolopidial receptor type tuned to 50-300 Hz vibrations, which projects into the central mechanosensory region specialised for processing low-frequency vibratory inputs, and most likely originates from the accessory organ or the posterior subgenual organ. Our findings contribute to understanding of the mechanical and neuronal basis of low-frequency vibration detection in insect legs and their highly differentiated sensory systems.

The mechanical leg response to vibration stimuli in cave crickets and implications for vibrosensory organ functions.

We investigate the influence of leg mechanics on the vibration input and function of vibrosensitive organs in the legs of the cave cricket Troglophilus neglectus, using laser Doppler vibrometry. By varying leg attachment, leg flexion, and body posture, we identify important influences on the amplitude and frequency parameters of transmitted vibrations. The legs respond best to relatively high-frequency vibration (200-2000 Hz), but in strong dependence on the leg position; the response peak shifts progressively over 500-1400 Hz towards higher frequencies following leg flexion. The response is amplified most strongly on the tibia, where specialised vibrosensory organs occur, and the response amplitude increases with the increasing frequency. Leg responses peaking at 800 and 1400 Hz closely resemble the tuning of the intermediate organ receptors in the proximal tibia of T. neglectus, which may be highly sensitive to positional change. The legs of free-standing animals with the abdomen touching the vibrating substrate show a secondary response peak below 150 Hz, induced by body vibration. Such responses may significantly increase the sensitivity of low-frequency receptors in the tibial accessory organ and the femoral chordotonal organ. The cave cricket legs appear suitable especially for detection of high-frequency vibration.

Independent suboesophageal neuronal innervation of the defense gland and longitudinal muscles in the stick insect (Peruphasma schultei) prothorax.

This study investigates the neuroanatomy of the defense gland and a related muscle in the stick insect Peruphasma schultei with axonal tracing and histological sections. The gland is innervated by three neurons through the Nervus anterior of the suboesophageal ganglion (SOG), the ipsilateral neuron (ILN), the contralateral neuron (CLN) and the prothoracic intersegmental neuron (PIN). The ILN has a large soma which is typical for motoneurons that cause fast contraction of large muscles and its dendrites are located in motor-sensory and sensory neuropile areas of the SOG. The CLN might be involved in the coordination of bilateral or unilateral discharge as its neurites are closely associated to the ILN of the contralateral gland. Close to the ejaculatory duct of the gland lies a dorsal longitudinal neck muscle, musculus pronoto-occipitalis (Idlm2), which is likely indirectly involved in gland discharge by controlling neck movements and, therefore, the direction of discharge. This muscle is innervated by three ventral median neurons (VMN). Thus, three neuron types (ILN, CLN, and PIN) innervate the gland muscle directly, and the VMNs could aid secretion indirectly. The cytoanatomy of motorneurons innervating the defense gland and neck muscle are discussed regarding the structure and functions of the neuropile in the SOG. As a basis for the neuroanatomical study on the defense gland we assembled a map of the SOG in Phasmatodea.

Four-dimensional light shaping: manipulating ultrafast spatiotemporal foci in space and time.

The spectral dispersion of ultrashort pulses allows the simultaneous focusing of light in both space and time, which creates so-called spatiotemporal foci. Such space-time coupling may be combined with the existing holographic techniques to give a further dimension of control when generating focal light fields. In the present study, it is shown that a phase-only hologram placed in the pupil plane of an objective and illuminated by a spatially chirped ultrashort pulse can be used to generate three-dimensional arrays of spatio-temporally focused spots. By exploiting the pulse front tilt generated at focus when applying simultaneous spatial and temporal focusing (SSTF), it is possible to overlap neighboring foci in time to create a smooth intensity distribution. The resulting light field displays a high level of axial confinement, with experimental demonstrations given through two-photon microscopy and the non-linear laser fabrication of glass.

The scolopidial accessory organs and Nebenorgans in orthopteroid insects: Comparative neuroanatomy, mechanosensory function, and evolutionary origin.

Scolopidial sensilla in insects often form large sensory organs involved in proprioception or exteroception. Here the knowledge on Nebenorgans and accessory organs, two organs consisting of scolopidial sensory cells, is summarised. These organs are present in some insects which are model organisms for the physiology of mechanosensory systems (cockroaches and tettigoniids). Recent comparative studies documented the accessory organ in several taxa of Orthoptera (including tettigoniids, cave crickets, Jerusalem crickets) and the Nebenorgan in related insects (Mantophasmatodea). The accessory organ or Nebenorgan is usually a small organ of 8-15 sensilla located in the posterior leg tibia of all leg pairs. The physiological properties of the accessory organs and Nebenorgans are so far largely unknown. Taking together neuroanatomical and electrophysiological data from disparate taxa, there is considerable evidence that the accessory organ and Nebenorgan are vibrosensitive. They thus complement the larger vibrosensitive subgenual organ in the tibia. This review summarises the comparative studies of these sensory organs, in particular the arguments and criteria for the homology of the accessory organ and Nebenorgan among orthopteroid insects. Different scenarios of repeated evolutionary origins or losses of these sensory organs are discussed. Neuroanatomy allows to distinguish individual sensory organs for analysis of sensory physiology, and to infer scenarios of sensory evolution.

The complex tibial organ of the New Zealand ground weta: sensory adaptations for vibrational signal detection.

In orthopteran insects, a complex tibial organ has evolved to detect substrate vibrations and/or airborne sound. Species of New Zealand weta (Anostostomatidae) with tympanal ears on the foreleg tibia use this organ to communicate by sound, while in atympanate species (which communicate by substrate drumming) the organ is unstudied. We investigated the complex tibial organ of the atympanate ground weta, Hemiandrus pallitarsis, for vibration detection adaptations. This system contains four sensory components (subgenual organ, intermediate organ, crista acustica homolog, accessory organ) in all legs, together with up to 90 scolopidial sensilla. Microcomputed tomography shows that the subgenual organ spans the hemolymph channel, with attachments suggesting that hemolymph oscillations displace the organ in a hinged-plate fashion. Subgenual sensilla are likely excited by substrate oscillations transmitted within the leg. Instead of the usual suspension within the middle of the tibial cavity, we show that the intermediate organ and crista acustica homolog comprise a cellular mass broadly attached to the anterior tibial wall. They likely detect cuticular vibrations, and not airborne sound. This atympanate complex tibial organ shows elaborate structural changes suggesting detection of vibrational stimuli by parallel input pathways, thus correlating well with the burrowing lifestyle and communication by substrate-transmitted vibration.

Neuronal Regression of Internal Leg Vibroreceptor Organs in a Cave-Dwelling Insect (Orthoptera: Rhaphidophoridae: Dolichopoda araneiformis).

Animals' adaptations to cave habitats generally include elaboration of extraoptic senses, and in insects the receptor structures located on the legs are supposed to become more prominent in response to constant darkness. The receptors for detecting substrate vibrations are often highly sensitive scolopidial sensilla localized within the legs or the body. For troglobitic insects the evolutionary changes in vibroreceptor organs have not been studied. Since rock is an extremely unfavorable medium for vibration transmission, selection on vibration receptors may be weakened in caves, and these sensory organs may undergo regressive evolution. We investigated the anatomy of the most elaborate internal vibration detection system in orthopteroid insects, the scolopidial subgenual organ complex in the cave cricket Dolichopoda araneiformis (Orthoptera: Ensifera: Rhaphidophoridae). This is a suitable model species which shows high levels of adaptation to cave life in terms of both phenotypic and life cycle characteristics. We compared our data with data on the anatomy and physiology of the subgenual organ complex from the related troglophilic species Troglophilus neglectus. In D. araneiformis, the subgenual organ complex contains three scolopidial organs: the subgenual organ, the intermediate organ, and the accessory organ. The presence of individual organs and their innervation pattern are identical to those found in T. neglectus, while the subgenual organ and the accessory organ of D. araneiformis contain about 50% fewer scolopidial sensilla than in T. neglectus. This suggests neuronal regression of these organs in D. araneiformis, which may reflect a relaxed selection pressure for vibration detection in caves. At the same time, a high level of overall neuroanatomical conservation of the intermediate organ in this species suggests persistence of the selection pressure maintaining this particular organ. While regressive evolution of chordotonal organs has been documented for insect auditory organs, this study shows for the first time that internal vibroreceptors can also be affected.

The scolopidial accessory organ in the Jerusalem cricket (Orthoptera: Stenopelmatidae).

Multiple mechanosensory organs form the subgenual organ complex in orthopteroid insects, located in the proximal tibia. In several Ensifera (Orthoptera), a small chordotonal organ, the so-called accessory organ, is the most posterior part of this sensory complex. In order to document the presence of this accessory organ among the Ensifera, the chordotonal sensilla and their innervation in the posterior tibia of two species of Jerusalem crickets (Stenopelmatidae: Stenopelmatus) is described. The sensory structures were stained by axonal tracing. Scolopidial sensilla occur in the posterior subgenual organ and the accessory organ in all leg pairs. The accessory organ contains 10-17 scolopidial sensilla. Both groups of sensilla are commonly spatially separated. However, in few cases neuronal fibres occurred between both organs. The two sensillum groups are considered as separate organs by the general spatial separation and innervation by different nerve branches. A functional role for mechanoreception is considered: since the accessory organ is located closely under the cuticle, sensilla may be suited to detect vibrations transferred over the leg's surface. This study extends the known taxa with an accessory organ, which occurs in several taxa of Ensifera. Comparative neuroanatomy thus suggests that the accessory organ may be conserved at least in Tettigoniidea.

Vibrational sensitivity of the subgenual organ complex in female Sipyloidea sipylus stick insects in different experimental paradigms of stimulus direction, leg attachment, and ablation of a connective tibial sense organ.

We document the sensitivity to sinusoidal vibrations for chordotonal organs in the stick insect tibia (Sipyloidea sipylus). In the tibia, the scolopidial subgenual organ (~40 scolopidial sensilla), distal organ (~20 scolopidial sensilla), and distal tibial chordotonal organ (~7 scolopidial sensilla) are present. We study the sensitivity of tibial sensory organs in all leg pairs to vibration stimuli as sensory thresholds by recording summed action potentials from Nervus cruris in the femur. The tibia was stimulated with a minishaker delivering vibrational stimuli. Because different experimental procedures may affect the vibration sensitivity, we here analysed possible effects of different experimental conditions: (1) the stimulus direction delivered in either horizontal or vertical direction to the leg; (2) recording responses only from the subgenual organ complex after ablation of the distal tibial chordotonal organ, and (3) the attachment of the leg to the minishaker by plastilin, beeswax-colophony, or freely standing legs. The tibial scolopidial organs give summed responses to vibration stimuli with highest sensitivity between 500 and 1000Hz for all leg pairs. In the different experimental series, we find that (1) thresholds were influenced by stimulation direction with lower thresholds in response to vertical vibrations, (2) ablating the distal tibial chordotonal organ by cutting the distal-most tibia did not change the summed sensory thresholds significantly, and (3) the attachment material between legs and the minishaker (plastilin or beeswax-colophony mixture) did not significant influence the sensory thresholds against free-standing tarsi. The distal tibial chordotonal organ is a connective chordotonal organ attached to a tendon and is likely a proprioceptive organ. These results emphasise that vibrational thresholds are mainly direction-sensitive. Thus, the direction of stimulus delivery during electrophysiological recordings is relevant for comparisons of vibratory sensory thresholds.

How many mechanosensory organs in the bushcricket leg? Neuroanatomy of the scolopidial accessory organ in Tettigoniidae (Insecta: Orthoptera).

The subgenual organ and associated scolopidial organs are well studied in Orthoptera and related taxa. In some insects, a small accessory organ or Nebenorgan is described posterior to the subgenual organ. In Tettigoniidae (Ensifera), the accessory organ has only been noted in one species though tibial sensory organs are well studied for neuroanatomy and physiology. Here, we use axonal tracing to analyse the posterior subgenual organ innervated by the main motor nerve. Investigating seven species from different groups of Tettigoniidae, we describe a small group of scolopidial sensilla (5-9 sensory neurons) which has features characteristic of the accessory organ: posterior tibial position, innervation by the main leg nerve rather than by the tympanal nerve, orientation of dendrites in proximal or ventro-proximal direction in the leg, and commonly association with a single campaniform sensillum. The neuroanatomy is highly similar between leg pairs. We show differences in the innervation in two species of the genus Poecilimon as compared to the other species. In Poecilimon, the sensilla of the accessory organ are innervated by one nerve branch together with the subgenual organ. The results suggest that the accessory organ is part of the sensory bauplan in the leg of Tettigoniidae and probably Ensifera.