Bimodal innervation of the infrared organ of Merimna atrata (Coleoptera,Buprestidae) by thermo- and mechanosensory units

The pyrophilous Australian “fire-beetle” Merimna atrata approaches forest fires and possesses abdominal infrared (IR) organs. Each round IR organ is centrally innervated by a sensory complex showing two different units: one thermoreceptive multipolar neuron and one mechanosensitive chordotonal organ (CO) consisting of two scolopidia. We investigated the CO and found that the scolopidia are mononematic (the scolopale cap remains below the cuticle) and monodynal (one sensory cell per scolopidium). The dendrites of the scolopidia extend anteriorly and are attached by their caps to the cuticle about in the middle of the absorbing area. Structural features at the site of innervation suggest that the CO measures minute thermal deformations caused by IR absorption. Therefore, an additional photomechanic component which has been described for the IR receptors of pyrophilous jewel beetles of the genus Melanophila can be proposed for the IR organ of Merimna. Because scolopidia can measure displacements in the subnanometer range, the CO may enhance the sensitivity of the IR organ. The sensory complex of the Merimna IR organ shows the same units and similar cuticular modifications as the tympanal organs of some noctuid moths. Therefore, a parallel evolution of insect ears and the Merimna IR organ is discussed.     

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.     

The Johnston's organ of three homopteran species: A comparative ultrastructural study

A transmission electron-microscopy study has been carried out on the pedicel of three homopteran species, with particular focus on the leafhopper Scaphoideus titanus Ball. The two other species, the planthoppers Hyalesthes obsoletus Signoret and Metcalfa pruinosa Say, were investigated in order to compare the ultrastructure of the Johnston's organ (JO) among representatives of the Auchenorrhyncha group. The results showed the presence of a well developed JO located within the pedicel. Depending on the species the JO is made of 25 up to 72 scolopidia arranged in a coronal array. Each scolopidium is connective, heterodynal, amphinematic and hosts three structurally dissimilar sensory neurons. Two of them have a type 1 ciliary segment while the third bears a type 2 cilium. The type 2 dendrite tip is associated with a tubular cap and is longer than the others, ending into the cuticle at the base of the flagellum. Other scolopidia with one or two neurons were found in S. titanus, forming an accessory organ. The presence of such a well developed mechanosensory apparatus is discussed in relation with the lifestyle of the three species.     

Johnston's organ in Neodiprion sertifer (Insecta: Hymenoptera)

The fine structure of Johnston's organ in the pine sawfly, Neodiprion sertifer, was studied by electron microscopy to determine if there exists a dimorphism in this organ corresponding to the sexual dimorphism in antennal shape and surface area. The organ is made up of scolopidia that are ultrastructurally similar to those of other insects. The scolopidia, identical in both sexes, comprise three sensory cells bearing two types of sensory processes: Two are shorter and smaller in diameter than the third, which extends into the cuticle of the membrane connecting pedicel and flagellum and terminates at an epicuticular invagination. The dendrites and sensory processes are surrounded by two types of enveloping (glial) cells-a scolopale cell and an attachment cell. Other enveloping cells occur at different levels of the scolopidium. Sexual dimorphism is evident only in the numbers of scolopidial groups: Males have more groups with fewer scolopidia, but both sexes possess about the same total number of scolopidia.     

Ultrastructure des organes chordotonaux des pièces céphaliques chez la larve du Speophyes lucidulus delar (coléoptère cavernicole de la sous-famille des bathysciinae)

Résumé Les organes chordotonaux présents dans les différentes pièces céphaliques de la larve du Speophyes peuvent être classés en deux catégories.La première catégorie regroupe les récepteurs scolopidiaux de l'antenne, du labium et du palpe maxillaire. On peut les comparer au scolopidium de l'organe tympanique du Criquet décrit par Gray (1960). La deuxième catégorie comprend les récepteurs scolopidiaux de la mandibule et de la lacinia: ils sont du amphinématique.Le sensille scolopidial de la galea représente un type intermédiaire.Nous signalons l'importance des structures de soutien et de fixation, qui doivent permettre une bonne transmission de toutes les déformations et tensions subies par le tégument. Nous discutons du rôle joué par la gap junction qui unit les deux dendrites dans les scolopidium. de la deuxième catégorie.Enfin nous essayons d'établir des hypothèses sur le fonctionnement des scolopidium.

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Fine structure of the chordotonal organs of the head appendages of Speophyes lucidulus larva

Summary The chordotonal organs located in the various head appendages of the Speophyes larva, can be divided into two classes.The scolopidial receptors of the antenna, the labium and the palpus maxillae belong to the first class. They can be compared to the scolopidium of the locust tympanic organ described by Gray.—The second class contains the scolopidial receptors of the mandible and the lacinia: their type is amphinematic.The scolopidial sensilla of the galea represents an intermediate type.We demonstrate many supporting and fixation structures which probably allow a good transmission of all the deformations and strains affecting the tegument.The function of the gap junction which connects the two dendrites in the scolopidia of the second class is discussed.Finally we try to formulate hypothesis of the functioning of scolopidia.

     

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.     

Differentiation of Apical and Basal Dendrites in Pyramidal Cells and Granule Cells in Dissociated Hippocampal Cultures

Hippocampal pyramidal cells and dentate granule cells develop morphologically distinct dendritic arbors, yet also share some common features. Both cell types form a long apical dendrite which extends from the apex of the cell soma, while short basal dendrites are developed only in pyramidal cells. Using quantitative morphometric analyses of mouse hippocampal cultures, we evaluated the differences in dendritic arborization patterns between pyramidal and granule cells. Furthermore, we observed and described the final apical dendrite determination during dendritic polarization by time-lapse imaging. Pyramidal and granule cells in culture exhibited similar dendritic patterns with a single principal dendrite and several minor dendrites so that the cell types were not readily distinguished by appearance. While basal dendrites in granule cells are normally degraded by adulthood in vivo, cultured granule cells retained their minor dendrites. Asymmetric growth of a single principal dendrite harboring the Golgi was observed in both cell types soon after the onset of dendritic growth. Time-lapse imaging revealed that up until the second week in culture, final principal dendrite designation was not stabilized, but was frequently replaced by other minor dendrites. Before dendritic polarity was stabilized, the Golgi moved dynamically within the soma and was repeatedly repositioned at newly emerging principal dendrites. Our results suggest that polarized growth of the apical dendrite is regulated by cell intrinsic programs, while regression of basal dendrites requires cue(s) from the extracellular environment in the dentate gyrus. The apical dendrite designation is determined from among multiple growing dendrites of young developing neurons.     

Light- and electron-microscopic analysis of a complex sensory organ: The tegula of Locusta migratoria

Summary Structure and organization of the tegula, a cupola-shaped structure located at the anterior base of the wings of locusts, is described using various morphological methods. Based on histological and cytological criteria, two different sensory systems are distinguished: (1) a field of mechanoreceptive hairs, and (2) a chordotonal organ. The total number of sensory cells corresponds to the number of axons within the nerve supporting the tegula. The hairs are situated at the posterior region of the tegula, and each hair is innervated by only one sensory cell. The complex architecture of the chordotonal organ is analyzed and the attachment of the scolopidia to the cuticle is described. A single scolopidium makes contact with several epidermal cells. The attachment cells run in parallel and are oriented longitudinally within the tegula, being connected to each other and to the epidermal cells by desmosomes. A function in relation to wing movements during flight is suggested for the two sensory systems within the mixed sense organ, tegula.     

Ultrastructure of the antennal sensilla of aphids

Summary An electron microscopical study of aphid antennal sensilla has revealed two types of trichoid sensilla. Type I, innervated by a single neuron is mechanoreceptive; type II, innervated by three to five neurons is both mechanoreceptive and chemoreceptive with possibly a third function. Johnston's organ in the pedicel comprises a peripheral ring of scolopidia inserted into the joint with the flagellum; two non-peripheral groups of scolopidia lie in the lumen with attachment points in the wall of the third segment. The fine structure of a campaniform sensillum on the pedicel is described together with two homologous and previously unknown sense organs at the joint between the fifth and sixth antennal segments. An unusually placed scolopidium in the lumen of the sixth segment has also been found. The function of this scolopidium is unknown but Johnston's organ, the campaniform sensillum and joint receptors are suggested to act as antennal proprioceptors.The authors thank the Long Ashton Research Station, Bristol for use of the SEM facilities. A.K. Bromley gratefully acknowledges the tenure of a S.R.C. CASE Studentship and thanks Professor L.H. Finlayson for research facilities     

Morphological differentiation of the embryonic peripheral neurons in Drosophila

Summary The stereotyped segmental and dorso-ventral organization of the peripheral nervous system (PNS) of Drosophila embryos allows the identification of all the neurons in the body wall. Distinct classes of neurons are distinguishable according to their location, the targets they innervate, the particular shape of their dendrites and their cell size. Those neurons innervating external sensory structures (es) and chordotonal organs (ch) have single dendrites and have been previously described (Ghysen et al. 1986; Dambly-Chaudiere and Ghysen 1986; Campos-Ortega and Hartenstein 1985). We describe here the identity and morphological features of three other classes of neurons in the body segments which have multiple dendrites (md neurons): 1) neurons that give rise to elaborate dendritic arborisations (da neurons); 2) neurons that have bipolar dendrites (bd neurons); 3) neurons that arborize around particular tracheal branches (td neurons). The thoracic hemisegment (T2 and T3) contains 13 da, one bd, one td, 21 es and four ch neurons; the abdominal hemisegment (A1 to A7) contains 14 da, three bd, three td, 15 es and eight ch neurons. The arrangement of the segmented peripheral neurons is highly invariant and provides a favorable assay system for the genetic analysis of neurodevelopment.     

The structure of a hair mechanoreceptor in the antennule of crayfish (Crustacea)

Summary In this study we examine the fine structure of mechanosensory hairs in the antennule of crayfish. The sensory hair is a stiff shaft with feather-like filaments. The hair's base is a large expansion of membrane which allows the hair shaft to deflect. The sensory transducing elements are located far from the hair, but are coupled mechanically with the hair shaft by a fine extracellular chorda. The sensory element is a type of scolopidium which consists of a scolopale cell and three sensory cells with a 9 + 0 type ciliary process.This type of scolopidium is characteristic of the chordotonal organ that has no cuticular structure on the surface of the exoskeleton. In this crustacean hair receptor, the deflection of the cuticular hair is transmitted through the chorda to the scolopidium which is a tension-sensitive transducer. The present study reveals that the mechanosensory hair of decapod crustaceans is a chordotonal organ accompanied by a cuticular hair structure. We also discuss comparative aspects of cuticular and subcuticular chordotonal organs in arthropods.     

Neural recognition molecules CHL1 and NB-3 regulate apical dendrite orientation in the neocortex via PTP alpha

Apical dendrites of pyramidal neurons in the neocortex have a stereotypic orientation that is important for neuronal function. Neural recognition molecule Close Homolog of L1 (CHL1) has been shown to regulate oriented growth of apical dendrites in the mouse caudal cortex. Here we show that CHL1 directly associates with NB-3, a member of the F3/contactin family of neural recognition molecules, and enhances its cell surface expression. Similar to CHL1, NB-3 exhibits high-caudal to low-rostral expression in the deep layer neurons of the neocortex. NB-3-deficient mice show abnormal apical dendrite projections of deep layer pyramidal neurons in the visual cortex. Both CHL1 and NB-3 interact with protein tyrosine phosphatase alpha (PTPalpha) and regulate its activity. Moreover, deep layer pyramidal neurons of PTPalpha-deficient mice develop misoriented, even inverted, apical dendrites. We propose a signaling complex in which PTPalpha mediates CHL1 and NB-3-regulated apical dendrite projection in the developing caudal cortex.     

The scolopidial organs in the first antennal segment in Allacma fusca (Collembola,Sminthuridae)

Summary The proprioceptors of the musculature moving the second antennal segment in the collembolan Allacma fusca were examined with the electron microscope. There are five muscles, the most important of which appear to be a levator and a depressor muscle. Only these two muscles are monitored by proprioceptors. These proprioceptors, the scolopidia, are referred to here as the levator (l-) and the depressor (d-) scolopidium. The former contains two, the latter one sensory cell.Both scolopidia deviate from the usual pattern in that they have unmodified dendritic outer segments (no dilatations, no electron-dense material) and are enveloped by only one cell. But l- and d-scolopidia also differ from one another. The dendritic inner segments of the l-scolopidium are branched to an h-shaped pattern, with one branch ending on the levator muscle and the other running to the antennal nerve where the perikarya are located. In the d-scolopidium a muscle fiber of about 1 m diameter (140–320 myosin filaments) accompanies the scolopidium for a distance of about 0.5 m.On the basis of the structural features it is hypothesized that (1) the mechanical forces possibly act on the membranes of the dendritic inner segments, (2) the small muscle parallel to the d-scolopidium is a receptor muscle, and (3) both scolopidia are highly derivative.Dedicated to Professor Dietrich Burkhardt on the occasion of his 60th birthday     

Embryogenesis of the connective chordotonal organ in the pedicel of the American cockroach: Cell lineage and morphological differentiation

Summary The ontogenesis of single scolopidia of the chordotonal organ of the American cockroach, Periplaneta americana, takes about 4 days. At 23% embryogenesis (100% = 30 d) the first anlagen of scolopidia were identified within the epithelium by staining with anti-horseradish peroxidase. Reconstruction of the cell lineage of the scolopidial cells was facilitated by two facts: (i) the arrangement of the cells throughout ontogenesis follows a strict pattern, and (ii) daughter cells are recognizable for several hours after mitosis by the cytoplasmic bridge and midbody joining them. When they separate, the midbody undergoes lysosomal degeneration in one of these cells. The earliest recognizable stage is a pair of cells, one of which (cell 1) encloses the other (cell 2) apically. The enclosing cell becomes the accessory cell. Cell 2 divides, yielding the mother cell (cell 3) of two sensory cells which degenerate later, and cell 2. Cell 2 gives rise to the attachment cell and to cell 2, which in turn produces the scolopale cell and the mother cell (cell 2 2) of a second pair of sensory cells; the latter are the definitive sensory cells. The end result is the total of 5 cells characteristic of the adult scolopidium. Secretion of the scolopale and cap together with the migration of the sensory cell perikarya into the antennal lumen complete development.     

Phosphorylation of CRMP2 is involved in proper bifurcation of the apical dendrite of hippocampal CA1 pyramidal neurons

The neural circuit in the hippocampus is important for higher brain functions. Dendrites of CA1 pyramidal neurons mainly receive input from the axons of CA3 pyramidal neurons in this neural circuit. A CA1 pyramidal neuron has a single apical dendrite and multiple basal dendrites. In wild‐type mice, most of CA1 pyramidal neurons extend a single trunk, or alternatively, the apical dendrite bifurcates into two daughter trunks at the stratum radiatum layer. We previously reported the proximal bifurcation phenotype in Sema3A?/?, p35?/?, and CRMP4?/? mice. Cdk5/p35 phosphorylates CRMP2 at Ser522, and inhibition of this phosphorylation suppressed Sema3A‐induced growth cone collapse. In this study, we analyzed the bifurcation points of the apical dendrites of hippocampal CA1 pyramidal neurons in CRMP2KI/KI mice in which the Cdk5/p35‐phosphorylation site Ser522 was mutated into an Ala residue. The proximal bifurcation phenotype was not observed in CRMP2KI/KI mice; however, severe proximal bifurcation of apical dendrites was found in CRMP2KI/KI;CRMP4?/? mice. Cultured hippocampal neurons from CRMP2KI/KI and CRMP2KI/KI;CRMP4?/? embryos showed an increased number of dendritic branching points compared to those from wild‐type embryos. Sema3A increased the number of branching points and the total length of dendrites in wild‐type hippocampal neurons, but these effects of Sema3A for dendrites were notobserved in CRMP2KI/KI and CRMP2KI/KI;CRMP4?/?hippocampal neurons. Binding of CRMP2 to tubulin increased in both CRMP2KI/KI and CRMP2KI/KI:CRMP4?/? brain lysates. These results suggest that CRMP2 and CRMP4 synergistically regulate dendritic development, and CRMP2 phosphorylation is critical for proper bifurcation of apical dendrite of CA1 pyramidal neurons. © 2012 Wiley Periodicals, Inc. Develop Neurobiol, 2013     

Polarized secretory trafficking directs cargo for asymmetric dendrite growth and morphogenesis

Proper growth of dendrites is critical to the formation of neuronal circuits, but the cellular machinery that directs the addition of membrane components to generate dendritic architecture remains obscure. Here, we demonstrate that post-Golgi membrane trafficking is polarized toward longer dendrites of hippocampal pyramidal neurons in vitro and toward apical dendrites in vivo. Small Golgi outposts partition selectively into longer dendrites and are excluded from axons. In dendrites, Golgi outposts concentrate at branchpoints where they engage in post-Golgi trafficking. Within the cell body, the Golgi apparatus orients toward the longest dendrite, and this Golgi polarity precedes asymmetric dendrite growth. Manipulations that selectively block post-Golgi trafficking halt dendrite growth in developing neurons and cause a shrinkage of dendrites in mature pyramidal neurons. Further, disruption of Golgi polarity produces neurons with symmetric dendritic arbors lacking a single longest principal dendrite. These results define a novel polarized organization of neuronal secretory trafficking and demonstrate a mechanistic link between directed membrane trafficking and asymmetric dendrite growth.     

The gustatory sensilla on the endophytic ovipositor of Odonata

The present paper aims at describing the fine structure of coeloconic sensilla located on the cutting valves of the endophytic ovipositor of two Odonata species, the anisopteran Aeshna cyanea (Aeshnidae) and the zygopteran Ischnura elegans (Coenagrionidae), by carrying out parallel investigations under SEM and TEM. In both species these coeloconic sensilla are innervated by four unbranched neurons forming four outer dendritic segments enveloped by the dendrite sheath. One dendrite terminates at the base of the peg forming a well developed tubular body, while the other three enter the peg after interruption of the dendrite sheath. The cuticle of the peg shows an apical pore and a joint membrane. This last feature, together with the tubular body and the suspension fibers, represent the mechanosensory components of the sensillum while the pore and the dendrites entering the peg allow chemoreception. The ultrastructural organization of these coeloconic sensilla is in agreement with the one reported for insect gustatory sensilla. Our investigation describes for the first time typical insect gustatory sensilla in Odonata. Electrophysiological and behavioral studies are needed to verify the role that these structures can perform in sensing the egg-laying substrata.     

Spindle-F Is the Central Mediator of Ik2 Kinase-Dependent Dendrite Pruning in Drosophila Sensory Neurons

During development, certain Drosophila sensory neurons undergo dendrite pruning that selectively eliminates their dendrites but leaves the axons intact. How these neurons regulate pruning activity in the dendrites remains unknown. Here, we identify a coiled-coil protein Spindle-F (Spn-F) that is required for dendrite pruning in Drosophila sensory neurons. Spn-F acts downstream of IKK-related kinase Ik2 in the same pathway for dendrite pruning. Spn-F exhibits a punctate pattern in larval neurons, whereas these Spn-F puncta become redistributed in pupal neurons, a step that is essential for dendrite pruning. The redistribution of Spn-F from puncta in pupal neurons requires the phosphorylation of Spn-F by Ik2 kinase to decrease Spn-F self-association, and depends on the function of microtubule motor dynein complex. Spn-F is a key component to link Ik2 kinase to dynein motor complex, and the formation of Ik2/Spn-F/dynein complex is critical for Spn-F redistribution and for dendrite pruning. Our findings reveal a novel regulatory mechanism for dendrite pruning achieved by temporal activation of Ik2 kinase and dynein-mediated redistribution of Ik2/Spn-F complex in neurons.     

Drosophila Hook-Related Protein (Girdin) Is Essential for Sensory Dendrite Formation

The dendrite of the sensory neuron is surrounded by support cells and is composed of two specialized compartments: the inner segment and the sensory cilium. How the sensory dendrite is formed and maintained is not well understood. Hook-related proteins (HkRP) like Girdin, DAPLE, and Gipie are actin-binding proteins, implicated in actin organization and in cell motility. Here, we show that the Drosophila melanogaster single member of the Hook-related protein family, Girdin, is essential for sensory dendrite formation and function. Mutations in girdin were identified during a screen for fly mutants with no mechanosensory function. Physiological, morphological, and ultrastructural studies of girdin mutant flies indicate that the mechanosensory neurons innervating external sensory organs (bristles) initially form a ciliated dendrite that degenerates shortly after, followed by the clustering of their cell bodies. Importantly, we observed that Girdin is expressed transiently during dendrite morphogenesis in three previously unidentified actin-based structures surrounding the inner segment tip and the sensory cilium. These actin structures are largely missing in girdin mutant. Defects in cilia are observed in other sensory organs such as those mediating olfaction and taste, suggesting that Girdin has a general role in forming sensory dendrites in Drosophila. These suggest that Girdin functions temporarily within the sensory organ and that this function is essential for the formation of the sensory dendrites via actin structures.