Two antagonistic functions of neural groups of the femoral chordotonal organ underlie thanatosis in the cricket Gryllus bimaculatus DeGeer

The cricket Gryllus bimaculatus displayed freezing (thanatosis) after struggling while the femoro-tibial joints of the walking legs were forcibly restrained. Myographic recording indicated that strong contraction of the flexor tibia muscle “leg flexion response” occurred under this restrained condition. During thanatosis, when the femoro-tibial joint was passively displaced and held for several seconds, it maintained its new position (catalepsy). Only discharge of the slow flexor units was mechanically indispensable for maintaining thanatosis and catalepsy. Differing roles of identified neuron subgroups of the femoral chordotonal organ were elucidated using this behavioral substrate. Ablation of the dorsal group neurons in the ventral scoloparium strengthened the leg flexion response and the normal resistance reflex, while ablation of the ventral group weakened both motor outputs. Ablation of the dorsal scoloparium neurons, or other main sensory nerves caused no detectable deficiency in femoro-tibial joint control. These results imply that both modes of flexor muscle activation promoted by the ventral group neurons are normally held under inhibitory control by the dorsal group. It is hypothesized that this antagonistic function causes immobilization of the femoro-tibial joint in a wide range of angles in thanatosis and catalepsy. Accepted: 12 November 1998     

Topographic mapping of the axons of the femoral chordotonal organ neurons in the cricket Gryllus bimaculatus

Central projections of the femoral chordotonal organ (FCO) neurons in the cricket Gryllus bimaculatus were investigated by selectively staining small numbers of axons. The FCOs in all legs consist of partly fused ventral and dorsal scoloparia in the proximal femur. The ventral scoloparium neurons can be reliably divided into two groups: the ventral group neurons (VG), which are arranged in a sequentially smaller manner distally, and dorsal group neurons (DG), which simply aggregate in the proximal region near the dorsal scoloparium. All axons of the FCO projected to the ipsilateral half of the respective thoracic ganglion. The VG axons possessed dorso-lateral branches in the motor association neuropile and antero-ventral branches dorso-lateral to the anterior ventral association centre. However, the more proximally the somata were situated, the more medially the main neurites terminated. The DG axons showed some variations: some axons of the distally located neurons possessed dorso-lateral branches and terminated on the boundary region of the mVAC, while the other axons terminated exclusively in the medical ventral association centre (mVAC), including the ventral part, which receives auditory sensory neuron projections. All axons of the dorsal scoloparium neurons projected exclusively into the dorsal part of the mVAC; however, the ventrally located neurons projected more ventrally than did the dorsally located neurons. The above characteristics were nearly identical in the pro- and metathoracic FCOs. These results suggest that the cricket FCO axons are roughly organized in a somatotopic map and are broadly differentiated in their function.     

The development of the sensory organs of the legs in the blowfly,Phormia regina

Summary The development of the sensory neurons of the legs of the blowfly,Phormia regina has been described from the third instar larva to the late pupa using immunohistochemical staining. The leg discs of the third instar larva contain 8 neurons of which 5 come to lie in the fifth tarsomere of the developing leg. Whereas 2 neurons persist at least to the late pupa, the other cells degenerate. The first neurons of gustatory sensilla arise in the fifth tarsomere at about 1.5 h after formation of the puparium. Most of these sensilla, however, appear within a short time period beginning at about 18 h. The femoral chordotonal sensory neurons first appear at the time of formation of the puparium, as a mass of cells situated in the distal femur. During later pupal development 2 groups of these cells come to lie at the femur-trochanter border, where they become the proximal femoral chordotonal organ of the adult; the remaining cells become the distal femoral chordotonal organ. Other scolopidial neurons appear later in development. The nerve pathways of the late pupal leg are established either by the axons of the cells that are present in the larval leg disc or by new outgrowing processes of sensory neurons. In the tibia, the initial direction of new outgrowth differs in different regions of the segment: proximal tibial neurons grow distally, while distal tibial neurons grow initially proximally.     

Selective expression of glionexin,a glial glycoprotein,in insect mechanoreceptors

The distribution of a glial cell-associated glycoprotein, glionexin (GX), on sensory receptors of the adult cricket Acheta domesticus is described, using the monoclonal antibody 5B12 as an immunohistochemical probe. GX was previously shown to be widely distributed in the embryo and to persist in the postembryonic to adult central nervous system. Here we demonstrate that it is restricted in the adult periphery to three subclasses of mechano-receptor sensilla: large socketed hair mechanoreceptors, their associated campaniform sensilla, and chordotonal organs. GX was not detected in photoreceptors, chemoreceptors, or other mechanoreceptors. The pattern of distribution differs significantly within the three subclasses of mechanoreceptors. In the hair and campaniform receptors GX is restricted to the extracellular space among glial cells clustered around the axon hillock region, but in chordotonal organs it surrounds the scolopidium at the tip of dendrites. The highly restricted distribution of GX in the periphery suggests possible functions that include mechanical stability of the sensory apparatus and ionic homeostasis in the respective neuronal spike-generating regions. The developmental modulation of GX expression is taken to imply multiple functions for the molecule during the life of the insect. 1994 John Wiley & Sons, Inc.     

Range fractionation in the locust metathoracic femoral chordotonal organ

Summary Insect femoral chordotonal organs are internal proprioceptors which monitor the position and movements of the femur-tibia joint of the leg. The locust (Locusta migratoria) metathoracic femoral chordotonal organ is composed of approximately 100 neurones with a variety of response properties. In this study intracellular recordings were used to examine the range fractionation of phasic and tonic responses to tibial movements. Some neurones responded across the full range of leg angles, while others had restricted response ranges, and could therefore act as labeled lines. Neurones with maximal firing at mid-angles are described for the first time in a locust femoral chordotonal organ. Responses are discussed in terms of underlying structural constraints on signal transduction.Abbreviation (mt) FCO (metathoracic) femoral chordotonal organ     

Mechanism of execution of sequential motor acts during copulation behavior in the male cricketGryllus bimaculatus DeGeer

Summary The roles of the mechanosensory afferents from the wings, cerci, tergites and genitalia in copulation behavior were examined by ablation and stimulation in the male cricketGryllus bimaculatus DeGeer.The sexually excited male cricket exhibited an intense posture (IP) upon contact stimulation of the elytra and the 4th to 9th abdominal tergites. This posture allowed the backward slipping (BWS) or hooking to take place subsequently.Backward slipping (BWS), which is the movement to get under the female, was elicited during IP by contact stimulation to the middle and distal regions of the dorsal surface of the cercus.Hooking, the coordinated movements for hanging the epiphallus onto the female's subgenital plate could be induced during IP by contact either on the dorsum, periproct or proximal 2 mm regions of the cercus. The latter two regions played a role in performing hooking accurately.Among four types of mechano-sensilla on the cercus the trichoid type was crucial for the initiation of BWS and hooking. Calculations revealed that about 60 trichoid hairs (4% of all the trichoid hairs in one cercus) were sufficient for the male to carry out hooking normally.The input from the bristle hairs on the epiphallus initiated the spermatophore extrusion (SPE) by swelling the endophallus.These results demonstrated that copulation behavior in the male cricket consisted of several motor acts and each act is triggered by specific input from the contact-sensitive sensilla on the elytra, tergites, cerci and genitalia. The sequential execution of each motor act is achieved because one motor act results in a positional change in contact with the female which in turn gives rise to another act. This type of motor control is a model of the so-called chain reaction in instinct behavior.Abbreviations BWS backward slipping - C cercus - EN endophallus - EP epiphallus - EPc epiphallic convexity - FW forewing - H hook - IP intense posture - HW hindwing - P pouch - PP periproct - SP spermatophore - SPE spermatophore extrusion - SEM scanning electron microscope     

An exaggerated trait in insects: The prothoracic skeleton of Stictocephala bisonia (Homoptera: Membracidae)

The prothoracic skeleton of Stictocephala bisonia was investigated in adults and fifth-instar nymphs on a gross morphological (SEM, maceration) and light microscopic level. In both nymphs and adults, the prothoracic skeleton consists of the pronotum, episternum, epimeron, precoxale, sternum, trochantin, and two endoskeletal characters (furcal arms and pleural apophyses). In nymphs, the entire pronotum is a single-layered outgrowth of the integument communicating with the body cavity and filled with hemolymph and fat body cells (“spine”); the dorsal and ventral processes and the suprahumeral bud are extensions of this single-layered integument. In adults, the pronotum is composed of (1) a proximal, single-layered part, and (2) a larger, distal, double-layered part (“posterior reduplication”) with two cuticular layers separated by a thin lumen. The posterior reduplication is elevated above the body and forms hollow (air-filled) extensions (e.g., suprahumeral horns). Its two cuticular layers are connected through cuticular columns that appear on the external surface as pits. The lumen between these layers communicates with the body cavity and contains nerves and tracheae. In the lumen of newly eclosed adults, intercellular space, epidermal cells with long processes, and hemocytes with nonlipid granules are present. In the lumen of sclerotized adult pronota, the intercellular space has disappeared, together with definite cell boundaries. Several structures are associated with the external cuticle: two types of innervated sensilla trichodea that articulate in the center of external pits, sensilla campaniformia, sensilla coeloconica, and cuticular canals with exterior openings. The morphogenetic implications of pronotal construction, various aspects of adult prothoracic anatomy, and the value of glands and sensilla for an adaptive interpretation of the pronotum are discussed. J. Morphol. 238:157–178, 1998. © 1998 Wiley-Liss, Inc.     

Adaptive reconfiguration of a reflex circuit during different motor programmes in the locust

The cuticle strain which develops in the hindleg tibiae when a locust prepares to kick, or when the tibia thrusts against an obstacle, is detected by two campaniform sensilla, which reflexly excite the fast extensor tibiae motoneuron, some of the flexor tibiae motoneurons and nonspiking interneurons. The reflex excitation is adaptive for the extensor motoneuron during both co-activation and thrusting, but is only adaptive for the flexor motoneurons during co-activation, and is maladaptive during thrusting. We show that the femoral chordotonal organ, which monitors tibial position, controls the efficacy of the strain feedback. The campaniform sensilla-induced depolarization in the extensor motoneuron is about twice as large when the tendon is in mid position (reflecting a tibial-femoral angle of 90°) than when fully stretched (reflecting tibial flexion), while in the flexors the reverse is true. The amplitudes of excitatory postsynaptic potentials evoked by single campaniform sensilla spikes, are, however, not affected. Our data suggests that the chordotonal organ modulates the gain of the strain feedback onto the motoneurons by exciting interneuronal circuits whose output sums with the former. Thrusting typically occurs with the tibia partially extended, therefore the actions of the chordotonal organ support the production of a maximal thrusting force. Accepted: 27 December 1996     

Differential Postalightment Oviposition Behavior of Monarch Butterflies on Asclepias Species

The monarch butterfly, Danaus plexippus L., oviposits mainly on plants in the Asclepiadaceae, particularly within the genus Asclepias. We studied postalightment oviposition behavior of monarch females on three host species—Asclepias curassavica, A. incarnata , and A. tuberosa. After landing on the host, they used their forelegs, midlegs, and antennae to assess plant suitability. When these appendages were examined by scanning electron microscopy, contact chemoreceptor sensilla were found. In choice tests, A. incarnata was most preferred, while A. tuberosa was least preferred. However, the use of appendages varied for the different host species. Antennae were most frequently used during post-alightment behavior on A. curassavica, whereas forelegs were used more often on A. incarnata, and all three appendages were used extensively on A. tuberosa. Use of the midlegs was generally followed by use of the antennae. Tasting with either forelegs or antennae apparently may lead to egg laying on some host species. Rupture of the plant surface by midleg spines was also observed. The behavior and host preference of individual females varied significantly and may reflect differences in receptor sensitivity.     

Ontogenèse de l'innervation des organes sensoriels des antennes chezReticulitermes lucifugus santonensis Feyt.

Summary The author has studied the ontogenesis of the innervation of neurosensorial organs on the antennæ ofReticulitermes lucifugus santonensis Feyt. She has found, among the organs with external processussensilla trichoidea (small, medium an big),sensilla basiconica, andsensilla campaniformia. Repartition and variation in number of sensilla during developpement have been described. She describe so 3 sensorial organs without any external processus: 2 chordotonal organs (one in the scapus, the other in the pedicellus). Antennal nerves and tracheas have been followed.

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Je tiens à exprimer toute ma reconnaissance à M. lePr Richard, pour m'avoir invitée dans son Laboratoire et pour l'intérêt qu'il a porté à mon travail Je remercie très vivement tous mes amis du Laboratoire de Biologie Animale qui m'ont aidée dans cette étude.     

Interaction between descending input and thoracic reflexes for joint coordination in cockroach: I. Descending influence on thoracic sensory reflexes

Tethered cockroaches turn from unilateral antennal contact using asymmetrical movements of mesothoracic (T2) legs (Mu and Ritzmann in J Comp Physiol A 191:1037–1054, 2005). During the turn, the leg on the inside of the turn (the inside T2 leg) has distinctly different motor patterns from those in straight walking. One possible neural mechanism for the transformation from walking to inside leg turning could be that the descending commands alter a few critical reflexes that start a cascade of physical changes in leg movement or posture, leading to further alterations. This hypothesis has two implications: first, the descending activities must be able to influence thoracic reflexes. Second, one should be able to initiate the turning motor pattern without descending signals by mimicking a point farther down in the reflex cascade. We addressed the first implication in this paper by experiments on chordotonal organ reflexes. The activity of depressor muscle (Ds) and slow extensor tibia muscle (SETi) was excited and inhibited by stretching and relaxing the femoral chordotonal organ. However, the Ds responses were altered after eliminating the descending activity, while the SETi responses remain similar. The inhibition to Ds activity by stretching the coxal chordotonal organ was also altered after eliminating the descending activity.     

Responses and locations of neurones in the locust metathoracic femoral chordotonal organ

Summary Insect legs possess chordotonal organs which monitor leg angle, and the direction, velocity and acceleration of leg movements. The locust metathoracic femoral chordotonal organ (mtFCO) has previously been studied morphologically and physiologically, but no detailed analysis of the responses of individual neurones, and their location in the organ has so far been produced. By recording from, and staining mtFCO neurones I have been able to compile for the first time such a map. The distribution of neurone somata in the locust mtFCO is more complex than previously thought: receptors sensitive to both stretch and relaxation of the apodeme are distributed throughout the organ. Seventeen response types were encountered. Neurones with a particular response type have somata in comparable locations within the mtFCO. Comparisons are made between the response types found in the stick insect and those in the locust. The possible functions of some of the responses are discussed.Abbreviation (mt)FCO (metathoracic) femoral chordotonal organ - F-T femur-tibia     

Sequential organization of grooming behaviors of the mosquito,Aedes triseriatus

Grooming behavior was studied in adult females of the mosquito Aedes triseriatus(Say). The grooming repertoire consisted of 12 different behaviors (accounting for bilateral symmetry) organized into five sequences. The proboscis and antennae were scraped by the forelegs, whereas the dorsal and ventral surfaces of the wings, and the forelegs, midlegs, hindlegs, and tip of the abdomen were scraped by the hindlegs. Each sequence ended with hindleg groomng. Tibial grooming combs were found on the ventral apices of the fore- and hind-tibiae but not on the mid-tibiae. A multidimensional scaling procedure grouped the grooming behaviors in two ways: (1) by the relative position of the groomed structure along the anteroposterior axis of the insect's body, and (2) by whether the groomed structure has a locomotory or sensory function. This suggests that mosquitoes groom both to clear sensilla of obstructive matter and to clean and smooth scales on legs and wings, possibly to decrease drag during flight.     

Sensory organs of the thoracic legs of the moth Manduca sexta

Summary The thoracic legs of the moth Manduca sexta acquire a new form and develop a new complement of sensory organs and muscles during metamorphosis from larva to adult. Because of our interest in the reorganization of neural circuitry and the acquisition of new behaviors during metamorphosis, we are characterizing sensory elements of larval and adult legs so that we may determine the contribution of new sensory inputs to the changes in behaviors. Here we describe the sensory structures of adult legs using scanning electron microscopy to view the external sensilla and cobalt staining to examine innervation by underlying sensory neurons. We find that, in contrast to larval legs, the adult legs are covered with a diverse array of sensilla. All three pairs of thoracic legs contain scattered, singly innervated scalelike sensilla. Campaniform sensilla occur singly or in clusters near joints. Hair plates, consisting of numerous singly innervated hairs, are also present near joints. Other more specialized sensilla occur on distal leg segments. These include singly innervated spines, two additional classes of singly innervated hairs, and three classes of multiply innervated sensilla. Internal sensory organs include chordotonal organs, subgenual organs, and multipolar joint receptors.     

昆虫弦音器及其声音感受分子机制

弦音器是昆虫类特有的一种机械感受器,亦称弦音感受器或剑梢感受器。它主要具有感知外界声压和体内肌肉运动的听觉功能,研究弦音器的机能结构对揭秘昆虫听觉的神经机制有重要的科学意义。本文从弦音器多样性和进化入手,重点综述了弦音器的微细结构、基因功能定位、声音感受分子机制及其声压增幅分子生物物理学原理,为昆虫听觉仿生学的研究提供了理论依据。     

Ontogenetic trajectories and early shape differentiation of treehopper pronota (Hemiptera: Membracidae)

Membracids (family: Membracidae), commonly known as treehoppers, are recognizable by their enlarged and often elaborated pronota. Much of the research investigating the development and evolution of this structure has focused on the fifth instar to adult transition, in which the pronotum undergoes the largest transformation as it takes on adult identity. However, little is known about the earlier nymphal stages, the degree to which the pronotum develops at these timepoints, and how development has changed relative to the ancestral state. Here, we studied the nymphal stages and adults of five morphologically distinct membracid species and of Aetalion reticulatum (family: Aetalionidae), the outgroup which was used as an ancestral state proxy. We found that shape differentiation in the pronotum of membracids can start as early as the second instar stage. Most shape differentiation occurs within the nymphal stages and not in the embryo since the shape of the first-instar pronotum did not differ from the outgroup species in all but one species we investigated. We found the anterior–posterior axis of the pronotum elongated at a faster relative rate in membracid species than in A. reticulatum, which contributed to the development of exaggerated pronotal size. Finally, we found differences in the morphogenesis of shape across species. We suggest this is due to the developmental and evolutionary divergence of differential growth patterning of the dorsal surface of the pronotum, not only across species, but also between stages within the same species. This lability may contribute to the evolvability and diversification of the membracid pronotum.     

Central projections of sensory cells of the midleg of the locust,Schistocerca gregaria

The central projections of trichoid hairs and of some scolopidial organs of the mesothoracic leg of the locust Schistocerca gregaria were studied by using nickel chloride backfilling and single cell recording. Trichoid hair sensilla on different parts of the legs project somatotopically in the ventral part of the ipsilateral neuropile of the mesothoracic ganglion. Generally, distally located receptors have their terminal arborizations in ventro-lateral areas of the neuropile, and proximally located receptors in ventro-medial areas. The axons of the subgenual organ and tarsal chordotonal organs project into the intermediate neuropile.     

Sensilla of Carabus (Isiocarabus) fiduciarius saishutoicus Csiki (Coleoptera : Carabidae)

Sensilla on the antenna of the ground beetle, Carabus (Isiocarabus) fiduciarius saishutoicus Csiki (Coleoptera: Carabidae), were investigated using scanning electron microscopy. In both sexes, 5 types of sensilla trichodea, 7 types of sensilla basiconica, 1 type of sensilla ampullacea, and 2 types of sensilla campaniformia are distinguished. No sexual differences in the types and distribution of sensilla are found on the antenna, except a ventral, depressed, hairless region of the male. Various types of sensilla present on the mouthparts, head, pronotum, elytron, hindwing, legs, abdominal tergites and sternites, aedeagus, and ovipositor are also described.     

L'ontogénèse des organes chordotonaux antennaires deCalotermes flavicollis (Fab.)

Summary In antennæ of Termites there are 3 groups of sense organs: — the chordotonal organ of the scapus;—the chordotonal organ of pedicellus; the Johnston's organ.Their structures are always the same during ontogenesis; the number of scolops increases slowly. Johnston's organ however distinguishes of the other in course of moulting but its modifications are very similar in structure to these of sensilla trichodea.

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Zusammenfassung Bei den Antennen der Termiten sind 3 Gruppen von Sinnesorganen feststellbar: das Chordotonal Organ des erstens Gliedes; das Chordotonal Organ des zweites Gliedes; das Johnston'sche Organ.Diese Struktur bleibt während der ganzen Ontogenese dieselbe; die Anzuhl der Scolopidialen Nägel steigt langsam an. Das Johnston'sche Organe hingegen insicht davon im Laufe der Häutungen ab, trotzdem seine Modificationen beim Häutungs denen der sensilla trichodea gleichen.

     

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.