Femoral chordotonal organ in the legs of an insect,Chrysoperla carnea(Neuroptera)

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 (9x2+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.     

Fine structure of the statocyst sensilla of the mysid shrimp Neomysis integer (Leach, 1814) (Crustacea,Mysidacea)

The fine structure of the statocyst sensilla of Neomysis integer was investigated. The statocyst contains about 35 sensilla, which are composed of two bipolar sensory cells, nine enveloping cells, and a seta. The sensory cells consist of an axon, a perikaryon, and a dendrite. The dendrite contains a proximal segment with a ciliary rootlet and at least one basal body, and a distal segment with a ciliary axoneme (9 × 2 + 0) at its base. The distal segment extends along the peripheral wall of the seta and is in close contact with the wall of the hair shaft. The enveloping cells surround the proximal and distal segments of the dendrite. The innermost enveloping cell contains a scolopale rod. It surrounds the receptor lymph cavity and secretes flocculent material into this cavity. From the tip of the cell a dendritic sheath, which encloses the distal segment of the dendrite, emerges. A peculiar feature of the second enveloping cell is the presence of a scolopale-like rod, which is more slender and less pronounced than in the first enveloping cell. The seta consists of three parts: a socket, a tubular midpart, and a gutter-like apical part, the tip of which penetrates into the statolith. The seta shows over its full length a bilaterally symmetrical axis that is coplanar with the plane in which the seta is bent toward the statolith. The structure of the seta and the position of the distal segments provide morphological evidence for directional sensitivity of the sensilla and for the magnitude of shear on the setal wall being an adequate stimulus.     

The metathoracic wing-hinge chordotonal organ of an atympanate moth,Actias luna (Lepidoptera,Saturniidae): a light- and electron-microscopic study

Summary The structure of a simple chordotonal organ, the presumed homologue of the noctuoid moth tympanal organ, is described in the atympanate moth, Actias luna. The organ consists of a proximal scolopidial region and a distal strand, which attaches peripherally to the membranous cuticle ventral to the hindwing alula. The strand is composed of elongate, microtubule-rich cells encased in an extracellular connective tissue sheath. The scolopidial region houses three mononematic, monodynal scolopidia, each comprised of a sensory cell, scolopale cell, and attachment cell. The dendritic apex is octagonally shaped in transverse section, its inner membrane lined by a laminated structure reminiscent of the noctuoid tympanal organ collar. A 9+0-type cilium emerges from the dendritic apex, passes through both the scolopale lumen and cap, and terminates in an extracellular space distal to the latter. Proximal extensions of the attachment cell and distal prolongations of the scolopale cell surrounding the cap are joined by an elaborate desmosome, with which is associated an extensive electron-dense fibrillar plaque. Within the scolopale cell, this plaque constitutes the scolopale rod material. The data are discussed in terms of both the organ's potential function, and its significance as the evolutionary proto-type of the noctuoid moth ear.     

Structure of the subgenual organ in the green lacewing, Chrysoperla carnea

REM and TEM studies of the subgenual organ in Chrysoperla carnea (Neuroptera: Chrysopidae) show that it is composed of three scolopidia, each with one sensory, one scolopale and one cap cell. The distal part of the dendrite shows a cilium with a '9 + 0' structure. The cross-handing pattern of the ciliary root has a periodicity of bands of about 61 nm. The scolopale material in a certain part of the scolopale cell is organized into five rods. The cell bodies of all three cap cells form a lens-like structure. the velum, which is fixed to the leg wall and the trachea with an extracellular material. The importance of the velum is discussed. Four types of intercellular junction are found; spot desmosomes. belt desmosomes, septate junctions and gap junctions.     

Structure of the tarso-pretarsal chordotonal organ in the imago of Tineola bisselliella Humm. (Lepidoptera : Tineidae)

The tarso-pretarsal chordotonal organ in the imago of Tineola bisselliella (Lepidoptera .: Tineidae) includes 2 groups of scolopidia. (1) A basal group composed of 2 scolopidia, each with one neuron and one scolopidium with 3 neurons, whose dendrites present a typical structure of cilium; the dendrite becomes dense apically; it is a common characteristic for all the mechanoreceptive dendrites. (2) An apical scolopidium with 3 neurons, whose dendrites have the same size and are covered with a conical cap.The combination of single and triple dendrites is unusual in the limb chordotonal organs. The 2 groups of scolopidia are not in contact. The importance of the support structures and fixation structures is discussed. In the ciliary root region, peculiar desmosomes occur between the dendrite and the scolopale cell. For the apical scolopidium, one attachment cell is seen distally and it appears in close association with the articular membrane.     

东亚飞蝗Locusta migratoria manilensis（Mey．）膝下器中具橛感器的附属细胞和附属结构的超微结构

东亚飞蝗膝下器的具橛感器主要由三类细胞组成．即：感觉细胞、感橛细胞和冠细胞。感觉细胞为具橛感器的主要结构和功能细胞,其超微结构已在其他的文章中描述。感橛细胞是具橛感器的主要支持细胞,从近端到远端依次与神经胶质细胞、感觉细胞的远端树突部分和感觉纤毛部以及顶端细胞外结构——冠、冠细胞直接接触．感橛细胞内最明显的结构为感概,另外,感橛细胞质被高度“空化”。冠细胞紧密包围着感橛细胞和冠,冠细胞中含有大量的纵行微管．并将整个具橛感器连接到体壁上。     

Sensilla on the maxillary palps of Helicoverpa armigera caterpillars: in search of the CO(2)-receptor

The ultrastructure of sensilla on the maxillary palps of helicoverpa armigera caterpillars has been investigated in order ot find candidates for CO(2)-receptors. The following sensilla are found on the palps: a) 8 chemosensory pegs at the tip; b), a large distal pore plate; c), a smaller proximal pore plate; d), a digitiform organ; e), a campaniform sensillum; and f), 3 scolopidia. Each chemosensory peg at the tip is innervated by 4-5 sensory neurons. Five of these pegs are most probably contact chemoreceptors, because each has a dendrite with a tubular body. The distal pore plate has a porous cuticle and is innervated by 3 sensory neurons, each of which sends a highly branched dendrite into a large cuticular cavity. The proximal pore plate is made up from two fused organs, has also a porous cuticle, and is innervated by two sensory neurons which send their dendrites into a narrow cuticular channel. The digitiform organ is innervated by one sensory cell which sends a highly lamellated dendrite into a narrow channel within a chip-shaped protrusion of the porous cuticle. For several reasons, the digitiform organ is the most probable candidate for the CO(2)-receptor. Another possible candidate is the distal pore plate.     

Axonal degeneration within the tympanal nerve of Schistocerca gregaria

This study describes time course and ultrastructural changes during axonal degeneration of different neurones within the tympanal nerve of the locust Schistocerca gregaria. The tympanal nerve innervates the tergit and pleurit of the first abdominal segment and contains the axons of both sensory and motor neurones. The majority of axons (approx. 97%) belong to several types of sensory neurones: mechano- and chemosensitive hair sensilla, multipolar neurones, campaniform sensilla and sensory cells of a scolopidial organ, the auditory organ. Axons of campaniform sensilla, of auditory sensory cells and of motor neurones are wrapped by glial cell processes. In contrast, the very small and numerous axons (diameter <1 microm) of multipolar neurones and hair sensilla are not separated individually by glia sheets. Distal parts of sensory and motor axons show different reactions to axotomy: 1 week after separation from their somata, distal parts of motor axons are invaded by glial cell processes. This results in fascicles of small axon bundles. In contrast, distal parts of most sensory axons degenerate rapidly after being lesioned. The time to onset of degeneration depends on distance from the lesion site and on the type of sensory neurone. In axons of auditory sensory neurones, ultrastructural signs of degeneration can be found as soon as 2 days after lesion. After complete lysis of distal parts of axons, glial cell processes invade the space formerly occupied by sensory axons. The rapid degeneration of distal auditory axon parts allows it to be excluded that they provide a structure that leads regenerating axons to their targets. Proximal parts of severed axons do not degenerate.     

Ultrastructure of a possible new type of crustacean cuticular strain receptor in Carcinus maenas (crustacea,decapoda)

A hitherto unknown sensillum type, the “intracuticular sensillum” was identified on the dactyls of the walking legs of the shore crab, Carcinus maenas. Each sensillum is innervated by two sensory cells with dendrites of “scolopidial” (type I) organization. The ciliary segment of the dendrite is 5–6 μm long and contains A-tubules with an electron-dense core and dynein arm-like protuberances; the terminal segment is characterized by densely packed microtubules. The outer dendritic segments pass through the endo- and exocuticle enclosed in a dendritic sheath and a cuticulax tube (canal), which is suspended inside a slit-shaped cavity by cuticular lamellae. The dendrites and the cavity terminate in a cupola-shaped invagination of the epicuticle. External cuticular structures are lacking. Three inner and four to six outer enveloping cells are associated with each intracuticular sensillum. The innermost enveloping cell contains a large scolopale that is connected to the ciliary rootlets inside the inner dendritic segments by desmosomes. Scolopale rods are present in enveloping cell 2. Since type I dendrites and a scolopale are regarded as modality-specific structures of mechanoreceptors, and since no supracuticular endorgan is present, the intracuticular sensilla likely are sensitive to cuticular strains. The intracuticular sensilla should be regarded as analogous to insect campaniform sensilla and arachnid slit sense organs.     

Functional morphology of the subgenual organ of the carpenter ant

Using light microscopy, confocal microscopy, electron microscopy and histochemistry, the subgenual organ (SGO) of an ant, Camponutas ligniperda, is investigated. Sensory units and attachment cells together enclose a large extracellular cavity, which is filled by acid mucopolysaccharides, as revealed by staining with ruthenium red. Due to this cavity, the whole SGO has the shape of a deformed sphere and the scolopidia exhibit a distribution of angles between 0 degrees and 60 degrees with the tibial long axis (as is shown by phalloidin-rhodamin staining of the actin filaments of the scolopale, viewed in situ by laser scanning confocal microscopy). The subgenual organ is innervated by a branch of the tibial nerve, which splits within or shortly distal to the femur-tibia joint. The other features of the SGO of Camponotus ligniperda are similar as in other insects: the SGO of Camponotus ligniperda contains about 35 scolopidial sensilla; it is fixed to the subgenual nerve on its proximal end, by its attachment cells to the opposite part of the cuticle; the fixation by the attachment cells is accomplished by a vast quantity of cytoplasmic microtubules; the construction of the sensory units is the same as in other mononematic scolopidial organs. The role of the extracellular lumen inside the organ and the special shape of the SGO of Camponotus ligniperda in mechanical transmission is discussed.     

Structure of the green lacewing tympanal organ (Chrysopa carnea,neuroptera)

Small swellings near the base of the radial vein in each fore wing of the green lacewing, Chrysopa carnea, resemble typical insect tympanal organs, but some important differences are apparent. The swellings are bounded dorsally and laterally by thick cuticle and ventrally by thin, membranous cuticle. The ventral membrane is formed by a single, thin sheet of exocuticle with flattened hypodermis internally, but lacks the tracheal component that forms part of the tympanum in the typical insect tympanal organ. The portion of the membrane beneath each swelling is rippled while proximally it is smooth. In contrast to typical insect tympanal organs, the swellings in C. carnea are largely fluid-filled since an unexpanded trachea runs through each organ. A distal and a proximal chordotonal organ composed of typical chordotonal sensory units are associated with each swelling. The distal organ contains from five to seven units while the proximal organ is composed of from 18 to 20 units. Each sensory unit is composed of three readily identifiable cells. Distally, an attachment cell unites with the membrane and is contiguous with the scolopale cell, which surrounds the dendrite of the bipolar neuron. On the basis of the morphological evidence, one would not expect these swellings to function as sound receptors. However, the results of physiological and behavioral experiments, presented elsewhere, show that these organs are receptors for ultrasound.     

Fine structural studies of Johnston's organ in the tobacco hornworm moth,Manduca sexta (Johannson)

There are approximately 650 scolopidial units in Johnston's organ of the tobacco hornworm moth, Manduca sexta. These sensory units exhibit greater complexity than noted previously (Gray, '60; Howse, '65; Uga and Kuwabara, '65; Moeck, '68; Ong, '69; Schmidt, '69). Each scolopale is innervated by a bipolar neuron whose dendrite terminals are modified into three ciliary structures. The largest of these differentiates near the tip into a multitubular structure. The remaining two cilia are structurally similar along the entire length of the scolopale shaft. From each of their bases, a collagen-like structure differentiates into numerous microtubules which extend proximally into individual channels of dendritic cytoplasm. A third channel, with a less developed root apparatus, was apparent for the largest cilium. Preliminary evidence suggests a proprioceptive function for this structure rather than an auditory one.     

The fine structure of haltere sensilla in the blowfly Calliphora erythrocephala (Meig.), with scanning electron microscopic observations on the haltere surface

The dipteran haltere incorporates large numbers of regularly disposed mechanoreceptors providing the sensory input enabling the vibrating haltere to function as a gyroscopic organ of equilibrium. Campaniform sensilla of the basal and scapal regions have been investigated by light and transmission electron microscopy, and these observations are augmented by scanning electron studies of the cuticle overlying the groups of sensilla. Each sensillum possesses a specialized fan-shaped terminal containing a complex and ordered association of microtubules and filaments. The transmission of stress to this region via the cuticle, and its possible role in transduction is considered. The fine structure of apical and basal sections of the distal sensory process and associated sheath cells is described; the functional significance of the distribution of mitochondria and other components is discussed. The organization of haltere chordotonal sensilla is described briefly, and compared with other mechanoreceptors with particular reference to microtubules and scolopale structures.     

Cytoskeletal elements in arthropod sensilla and mammalian photoreceptors.

Ciliary receptor cells, typified by cilia or modified cilia, are very common in the animal kingdom. In addition to the cytoskeleton of their ciliary processes these receptors possess other specific prominent cytoskeletal elements. Two representative systems are presented: i) scolopidia, mechanosensitive sensilla of various arthropod species; and ii) photoreceptor cells of the retina of the bovine eye. Two cytoskeletal structures are characteristic for arthropod scolopidia: a scolopale typifies the innermost auxiliary cell, and long ciliary rootlets are extending well into the sensory cells. The latter element is also characteristic for the inner segment of the photoreceptor cells in bovine. The scolopale of scolopidia is mainly composed of actin filaments. In the absence of myosin, the uniform polarity of the actin filaments and their association with tropomyosin all indicate a stabilizing role of the filament bundles within the scolopale. This function and a certain elasticity of actin filament bundles may be important during stimulation of the sensilla. The ciliary rootlets of both systems originate at the basal bodies at the ciliary base of the sensory cells and project proximally. These rootlets are composed of longitudinally oriented, fine filaments forming a characteristic regular cross-striation. An alpha-actinin immunoreactivity was detected within the ciliary rootlets of scolopidia. In addition, antibodies to centrin react with the rootlets of both types of receptors. Since centrin is largely responsible for the contraction of the flagellar rootlets in green algae, contraction may also occur in the ciliary rootlets of insect sensilla and vertebrate photoreceptors. In both systems, contraction or relaxation of the ciliary rootlets could serve in sensory transduction or adaptation.     

Are the funnel-canal organs the ‘campaniform sensilla’ of the shore crab,Carcinus maenas (Decapoda,Crustacea)?

Summary The funnel-canal organs on the dactyls of the shore crab, Carcinus maenas, are innervated by 3–24 sensory cells with unbranched dendrites, which attain a length of 500–1400 m. The outer dendritic segments are enclosed in a dendritic sheath and pass through the cuticle within a canal. Two dendrite types can be distinguished according to ultrastructural criteria: Type I has a long ciliary segment, A-tubules with an osmiophilic core and arms, and a thick ciliary rootlet. Type II possesses only a short ciliary segment and a thin ciliary rootlet. Each funnel-canal organ contains two type-I dendrites. Their ciliary bases appear a few m distal to those of the type-II dendrites (1 to 22 in number). Two inner and two to eight outer enveloping cells belong to a sensillum. The innermost enveloping cell contains a large scolopale. In the second enveloping cell single scolopale rods are present. Thus, the funnel-canal organs are characterized by structural features typical for mechano-sensitive scolopidia, on the one hand, and for chemoreceptors, on the other. Therefore, the funnel-canal organs are very likely bimodal sensilla (contact chemoreceptors). A comparison with other arthropod sensilla shows that cuticular mechanoreceptors of aquatic crustaceans generally exhibit a scolopidial organization.     

Structure of the abdominal receptor responsive to internally applied pressure in the blood-feeding insect,Rhodnius prolixus

Summary The sensory receptor responsive to pressure applied internally to the ventral abdominal body wall of the blood-feeding insects, Rhodnius prolixus, is a single sense cell containing, at its distal end, a cilium enclosed within a scolopale, a densely staining structure characteristic of insect scolopidial sensilla. A small spherical structure lies within a dilation near the midregion of the cilium, and contains nine heavily staining bodies, the position of each corresponding to a pair of microtubules in the cilium. Proximal to the dilation, the microtubules are organized in a ring of nine pairs with one microtubule of each pair associated with dyneinlike arms. Dastal to the dilation a central pair of microtubules is present, but dyneinlike arms are absent. The scolopale cell, which gives risc to the scolopale, has cytoplasmic invaginations that form an elaborate array of extracellular compartments surrounding the body wall of the sense cell. These compartments may serve to dampen high frequency vibrations permitting the receptor to respond to pressure exerted by touch, an attribute in keeping with the receptor's proposed function of detecting abdominal distension related to the size and movement of the stomach.     

Ultrastructure and blood–nerve barrier of chordotonal organs in the Drosophila embryo

Chordotonal organs of Drosophila embryos have become models for studies of developmental biology and molecular genetics due to their consistent segmental placement and mutability. Our first goal was to find the origin and anatomical correlate of the blood–nerve barrier of this PNS proprioreceptor in wild type embryos. The concept of a blood–nerve barrier for the PNS of the Drosophila embryo is new, and the present data are the first in this regard. A second goal was to reveal the ultrastructure of these four-celled stretch receptors, focusing particularly on the ‘core’ of this organ: the bipolar neuron enclosed by a scolopale cell. These latter data have resulted in a graphic reconstruction of the chordotonal organ which reveals how the four consistent cells fit together. At Stage 13 we first observed a clearly recognizable scolopale cell with an enclosed neuron. Surprisingly, an operative blood–nerve barrier, comprised of occlusive pleated-sheet septate junctions, exists at this relatively early stage. A blood–brain barrier is not yet functioning in the CNS during this same stage, as the perineurium is not present until Stage 17. Cross-sectional views of a more mature chordotonal organ show that the neuron’s inner segment has a ‘tongue-in-groove’ formation which fits the dendrite into the scolopale cell. Other newly discovered fine structural features are: hemidesmosomes linking individual scolopale rod bundles to the inner dendrite, and a cap cell matrix bonding with the tip of the ciliary dendrite. Functional aspects of these findings are discussed.     

Fine structure of scolopidia in Johnston's organ of female Aedes aegypti compared with that of the male.

The Johnston's organ of the female mosquito, Aedes aegypti, has only three types of scolopidia: types A, B, and C. It lacks the type D scolopidium of the male's organ. The basic structure and the location of each type in the female are similar to the counterparts in the male's organ. A single scolopidium is composed of a scolopale cell, an envelope cell, a long cap, and a third sheath, in addition to the two electron-dense scolopales produced inside the cytoplasm of two satellite cells. Each scolopidium has either two (type A) or three (type B) sensory cells. A type C scolopidium, mononematic in contrast to the amphinematic types A and B scolopidia, has two sensory cells, a microtubular cap cell, two microtubular accessory cells, and a scolopale cell with an intracellular scolopale. Even though the female Johnston's organ has all the components of the male's organ except for the single type D scolopidium, the female's organ shows relatively poorer organization and development. The female has a smaller and thinner basal plate, shorter and thicker prongs, fewer type A sensory cells, and a shorter flagellar flange, in addition to the overall smaller size of the pedicel.The probable function of each scolopidial type is discussed, especially in connexion with the probable identification of a single auditory sensillum in the male.     

The hair-peg organs of the shore crab,Carcinus maenas (Crustacea,Decapoda): Ultrastructure and functional properties of sensilla sensitive to changes in seawater concentration

Summary The hair-peg organs of the shore crab, Carcinus maenas, are modified hair-sensilla. A small hair shaft (peg) is surrounded by a tuft of solid cuticular bristles (hairs). Each hair-peg organ is innervated by 6 sensory neurons, 2 of which have scolopidial (type-I) dendrites. The outer segments of all dendrites pass through a cuticular canal extending to the articulated hair base in which the 2 type-I dendrites terminate. The other 4 (type-II) dendrites reach the clavate tip of the hair shaft and have access to a terminal pore and a large sickle-shaped aperture. Three inner and 8–12 outer enveloping cells belong to a hair-peg organ. The innermost enveloping cell contains a scolopale, which has desmosomal connections to the ciliary rootlets of the type-I dendrites. An inner and an outer sensillum lymph space are present. The ultrastructural features of the dendrites and the cuticular apparatus indicate that the hair-peg organs are bimodal sensilla, comprising 2 mechano- and 4 chemosensitive sensory neurons. Extracellular recordings from the leg nerve indicate that the chemosensitive neurons of the hair-peg organs respond to changes in seawater concentration in the physiological range of Carcinus maenas.Supported by the Deutsche Forschungsgemeinschaft (SFB 45/A1; W. Gnatzy)     

Sensory cell degeneration in the ontogeny of the chemosensitive sensilla in the labial palp-pit organ of the butterfly,Pieris rapae L. (Insecta,Lepidoptera)

Summary The ontogeny of the chemoreceptive sensilla in the labial palp-pit organ was studied in Pieris rapae by examining twelve successive stages between pupation and emergence of the imago, which takes a period of 160 h under the experimental conditions. Mitoses occur until 20 h after pupation. They lead to anlagen of sensilla, 91% of which are comprised of three sensory cells. However, two sensory cells degenerate in each sensillum during a period of 28 h. The same process occurs in anlagen with four sensory cells resulting in bicellular sensilla. Axons grow out only after the number of sensory cells has been reduced. Further consecutive steps in sensory cell differentiation are: (a) outgrowth of dendritic outer segment and dendrite sheath; (b) outgrowth of trichogen process and change in structure of elongating dendrite sheath; (c) deposition of cuticle and pore tubules in the pegs; (d) retraction of trichogen process; (e) increase in diameter of dendritic outer segment accompanied by increase of microtubule number and appearance of regularly spaced electron-dense bodies at tubular doublets; (f) branching of dendritic outer segment; and (g) transformation of the dendritic branches into curled lamellae and partial destruction of the dendrite sheath. The unique process of sensory cell degeneration is interpreted as an event that revokes a step towards a possible functional improvement of the labial palp-pit organ during further evolutionSupported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)