Structure of sensilla, olfactory perception, and behaviour of the bedbug, Cimex lectularius, in response to its alarm pheromone

When deprived of the terminal antennal segments, male and female bedbugs failed to respond to their alarm pheromone and to their assembling scent. Trans-oct-2-en-1-al or trans-hex-2-en-1-al, being the major constituents of the former, induce in adults and larvae of Cimex lectularius a typical alarm behaviour resulting in dispersal of assembled bedbugs; the rapidity of escape depends on the aldehyde concentration in the air. The behavioural threshold for adults is about 9×10¹⁴ molecules of trans-oct-2-en-1-al or 6×10¹⁵ molecules of trans-hex-2-en-1-al per ml air.The distal part of the terminal antennal segment of C. lectularius reveals the following sensilla: bristles (type A1), immersed cones (type B1), plates (type B2), grooved pegs (type C), smooth pegs (type D), hairs with even (type E1), and uneven wall thickness (type E2). The number and distribution of these sensilla is relatively constant and similar in both sexes, but differs slightly in neonate larvae. The pegs and hairs of types C, D, E1 and E2 were shown to have porous walls, a prerequisite for olfactory function.Receptor potentials were recorded from olfactory sensilla of types E1 and E2 after stimulation with trans-hex-2-en-1-al and trans-oct-2-en-1-al. The minimal concentration of trans-hex-2-en-1-al evoking a receptor potential is about 2×10¹⁰ molecules per ml air. The above olfactory sensilla were found to respond also to hexan-1-al, but almost no responses to pentan-1-al, butan-1-al, trans-hex-2-ene, and trans-oct-2-ene were observed. A minimum chain length of six carbons atoms and a terminal carbonyl group are molecular prerequisites for optimal odorant activity, while the presence of a Δ²-double bond is not essential for stimulation of the alarm pheromone receptors of the bedbug.     

Occurrence of certain cuticular structures confirms functionality of dorsal abdominal scent glands in Acanthosomatidae (Heteroptera: Pentatomoidea)

Elasmucha ferrugata (Fabricius, 1787) (Heteroptera: Acanthosomatidae) provides maternal care of eggs and larvae. Adults of both sexes have functional anterior dorsal abdominal scent glands (DAGs). Study of all internal and external cuticular structures of DAGs revealed that no DAGs are fully functional in the 1st larval instar, and very probably they are only slightly functional in the 2nd instar. Median and posterior DAGs are undoubtedly not functional in adults. There exists sexual dimorphism in the number of multicellular glandular units in anterior glands in adults. The occurrence of cuticular ductules of these units proves these to be functional glands. This is best considered in combination with the findings of a well-developed evaporatorium. Developed cuticular intima of the gland reservoir and/or the nearly closed ostiole or ostiolar scar bears no information about the functionality of the gland.     

Compartments in the abdomen of Drosophila and the role of the engrailed locus

A clonal analysis has shown that the dorsal surface of the first abdominal segment of Drosophila melanogaster is subdivided into anterior and posterior compartments. Cells of the posterior compartment grow up to but not beyond the anterior-posterior compartment border within the first abdominal segment and the intersegmental border that defines the boundary between the first and second abdominal segments. Growing within these boundaries, a narrow band of tissue clonally isolated from the adjoining tissue is formed. When these posterior cells are deficient for the engrailed locus, however, neither the compartment nor the segment border is maintained. The implications, that compartmentalization is essential for segmentation, and that all insect segments are subdivided by anterior and posterior compartments, are discussed.     

The comparative morphology of epidermal glands in Pentatomoidea (Heteroptera)

The Heteroptera show a diversity of glands associated with the epidermis. They have multiple roles including the production of noxious scents. Here, we examine the cellular arrangement and cytoskeletal components of the scent glands of pentatomoid Heteroptera in three families, Pentatomidae (stink bugs), Tessaratomidae, and Scutelleridae (shield-backed bugs or jewel bugs). The glands are; (1) the dorsal abdominal glands, (2) the tubular glands of the composite metathoracic gland, and (3) the accessory gland component of the composite metathoracic gland. The dorsal abdominal glands are at their largest in nymphs and decrease in size in adults. The metathoracic gland is an adult-specific gland unit with a reservoir and multiple types of gland cells. The accessory gland is composed of many unicellular glands concentrated in a sinuous line across the reservoir wall. The lateral tubular gland is composed of two-cell units. The dorsal abdominal glands of nymphs are composed of three-cell units with a prominent cuticular component derived from the saccule cell sitting between the duct and receiving canal. The cuticular components that channel secretion from the microvilli of the secretory cell to the exterior differ in the three gland types. The significance of the numbers of cells comprising gland units is related to the role of cells in regenerating the cuticular components of the glands at moulting in nymphs.     

Fine structure of the sensory organs of Drosophila melanogaster Meigen larva (Diptera : Drosophilidae)

Fine structure of the prominent external and internal sensory organs of Drosophila melaogaster Meigan (Diptera : Drosophilidae) larva was determined by transmission electron microscopy (TEM). The external sensory organs, namely, antennal, maxillary, ventral and labial organs, dorsal pits, sensory cones on the 8th and 9th abdominal segments, and the sensory hairs on the body of the larva, were studied. A new knob in pit (KIP) sensillum innervated by 3 dendrites was found on the dorsolateral surface on either side. Four tufts of hairs at the posterior end of each great lateral tracheal trunk were found to be sensory with a dendrite at the base of each tuft.Internally in the pharynx of the larva, 3 groups of symmetrically located sensory organs, namely, the anteroventral, dorsal, and posteroventral groups were found. In all they contain 10 pairs of sensilla. Amongst them, 2 pairs of sensilla were found to be of the compound type; one having 9 dendrites arranged in 3 groups of 4, 3 and 2; while the other had 6 dendrites grouped as 2 and 4. In addition, 2 groups of sensilla were found on the internal dorsal fold on either side.Similarities were observed in the dendritic organisation of sensilla in the internal mouthparts of the Drosophila larva and the adult. Unlike nerves of the adult, the larval nerves connected with the dorsal and maxillary organs have a significantly thick layer of glial cells ensheathing the bundle of axons.     

Morphology and Volatile Compounds of Metathoracic Scent Gland in Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae)

Tessaratoma papillosa (Drury) (Hemiptera: Tessaratomidae) is a serious insect pest of litchi and longan in South China. When disturbed, this insect could release large quantities of disagreeable odorous volatiles from its scent gland. Knowledge on the scent gland and its secretion is crucial for developing the semiochemical methods to manage this pest. Morphology and ultrastructure of the metathoracic scent glands (MTGs) were studied under stereo and scanning electron microscopy, and the volatile compounds of MTGs from both male and female T. papillosa were analyzed with coupled gas chromatography?Cmass spectrometry (GC?CMS). The MTG complex is located between the metathorax and the first abdominal segment at the ventral surface of the insect, which has a well-developed single double valve cystic-shaped orange median reservoir, paired colorless lateral glands in both sides, and a long and wavy tubular accessory gland that inlays tightly into the ventral edge around the median reservoir. The MTG opens to the body surface through paired ostioles located between the meso- and metacoxae of the evaporatorium with mushroom bodies. The GC?CMS analyses showed that female and male adults have nine major volatile components in common. Tridecane is the most abundant in both females and males, reaching up to 47.1% and 51.8% of relative amount, respectively. The minor component is benzophenone with only 0.28% and 0.14%. Furthermore, undecane, tetradecane, 3-methyl-tridecane, and cyclopentadecane were found only in males. The possible function of volatile compounds of MTG contents in T. papillosa is addressed.     

Stomodeal and neurohypophysial placodes in Ciona intestinalis: insights into the origin of the pituitary gland

The ascidian larva has a central nervous system which shares basic characteristics with craniates, such as tripartite organisation and many developmental genes. One difference, at metamorphosis, is that this chordate-like nervous system regresses and the adult's neural complex, composed of the cerebral ganglion and associated neural gland, forms. It is known that neural complex differentiation involves two ectodermal structures, the neurohypophysial duct, derived from the embryonic neural tube, and the stomodeum, i.e. the rudiment of the oral siphon; nevertheless, their precise role remains to be clarified. We have shown that in Ciona intestinalis, the neural complex primordium is the neurohypophysial duct, which in the early larva is a short tube, blind anteriorly, with its lumen in continuity with that of the central nervous system, i.e. the sensory vesicle. The tube grows forwards and fuses with the posterior wall of the stomodeum, a dorsal ectodermal invagination of the larva. The duct then loses posterior communication with the sensory vesicle and begins to grow on the roof of the vesicle itself. The neurohypophysial duct differentiates into the neural gland rudiment; its dorsal wall begins to proliferate neuroblasts, which migrate and converge to build up the cerebral ganglion. The most anterior part of the neural gland organizes into the ciliated duct and funnel, whereas the most posterior part elongates and gives rise to the dorsal strand. The hypothesis that the neurohypophysial duct/stomodeum complex possesses cell populations homologous to the craniate olfactory and adenohypophysial placodes and hypothalamus is discussed.     

FMRfamide-like immunoreactivity in the ventral nerve cord of the larval eastern spruce budworm,Choristoneura fumiferana (clemens) (Lepidoptera : Tortricidae)

Distribution of FMRFamide-like immunoreactivity was examined in the larval ventral nerve cord of the eastern spruce budworm, Choristoneura fumiferana (Lepidoptera : Tortricidae). Indirect immunofluorescent methods revealed the existence of 3 groups of FLI neurons in each ganglion. The neurons are distributed in a bilaterally symmetrical fashion at the anterodorsal, midlateral and posteroventral regions of the ganglia. There are 4 FMRFamide-like immunoreactive fiber tracts on the dorsal surface of the ganglia to which the anterodorsal FLI neurons project ipsilaterally, while the midlateral pair projects both ipsi-, and contralaterally. The last abdominal ganglion (AG8) has 4 additional pairs of FLI neurons; and axons from some of these extend into the median abdominal nerve, which suggests some role for this neuropeptide in the control of posterior structures of the larva.     

Respiratory system of the primitive moth Micropterix calthella (Linnaeus) (Lepidoptera : Micropterigidae)

The 1st thoracic spiracular atrium is closed by anterior and posterior muscle fibres extending between its dorsal and ventral wall. The 2nd thoracic spiracle has only a single (anterior) closing lip, movable by a muscle inserting on the wall below the spiracular aperture; this configuration may be a lepidopteran ground-plan autapomorphy. There are functional spiracles on abdominal segments I – VII, each with a closing “bow” and “lever”. There are intrinsic occlusor muscles in all abdominal spiracles and the 1st spiracle has an extrinsic (ventral) dilator. Dorsal dilator muscles or ligaments are absent. A dorsal and a ventral tracheal trunk extend from the 1st thoracic spiracle into the head; the latter supplies the mouthparts and the antenna; there is no connection between the dorsal and ventral cephalic trunk systems. There is a single series of lateral connectives between the spiracles of each side. There is a ventral tracheal commissure in both pterothoracic segments, but none in the prothorax. In each pterothoracic segment an anterior and a posterior tracheal arch give off branches to the wing and anastomose with each other on their downwards course into the leg. Wing tracheation is greatly reduced. The anterior and posterior tracheae of each wing are independent of each other. There is a dorsal commissure in abdominal segment VIII; ventral abdominal commissures are lacking in Micropterix, although present in other micropterigid genera. The terminalia are partly supplied from tracheae arising in segment VII. Air sacs occur in the tibiae only. Phylogenetic aspects of holometabolan tracheation patterns are discussed.     

Scanning electron microscopy of Drosophila melanogaster embryogenesis: III. Formation of the head and caudal segments

The formation of both the anterior most and posterior most segments in higher dipteran embryos involves complex movements of primordia which can be best visualized with the scanning electron microscope. During head formation, the gnathocephalic segments partially involute through the stomodeum. The labial segment forms the floor of the mouth, and the fused maxillary and mandibular segments form the lateral sides of the mouth. The involuted clypeolabrum forms the roof of the mouth. Invaginations of cells for segmentally derived sense organs can be found prior to involution on all the gnathocephalic and thoracic segments as well as on the labrum. The antennal sense organ derives from the lateral surface of the procephalic lobe. Following involution of the mouth parts, the dorsal ridge, which arises just anterior to the first thoracic segment, is drawn over the dorsal procephalic lobe producing the deep dorsal sac. The optic lobes of the brain invaginate anterior to the dorsal ridge just prior to the covering over of the head. The formation of the anal segment is similarly complex. Two rudimentary segments are found posterior to the eighth abdominal segment. During shortening of the germ band, the posterior most segment is drawn around the posterior tip of the embryo to lie ventrally. Two large anal pads form lateral to the anus from this segment. The next segment, following dorsal closure, produces a pair of anal sense organs and a central tuft of setae. Finally, the eighth abdominal segment gives rise to the posterior spiracles. Following dorsal closure these three segments fuse to produce the terminal (anal) segment of the larva.     

Biologically active abietane and ent-kaurane diterpenoids and other constituents from Erythroxylum suberosum

Bioassay-guided fractionation of the ethanol extract from the branches of Erythroxylum suberosum, which was toxic to brine shrimp larvae, afforded five diterpenes bearing abietane and ent-kaurane-type skeletons from an active fraction. From these, four were new, 7-oxo-16-hydroxy-abiet-15(17)-en-19-al, 16-hydroxyabiet-15(17)-en-7-one, 7α,16-dihydroxy-abiet-15(17)-en-19-al and ent-12α-hydroxy-kaur-16-en-19-al, while methyl ent-7α,15β-dihydroxy-kaur-16-en-19-oate is reported for the first time as a natural product. This is also the first reported occurrence of abietane-type diterpenes in the genus Erythroxylum. The flavonoid ombuin-3-rutinoside was isolated from an inactive fraction, while rutin (quercetin-3-rutinoside) was obtained from the non-toxic ethanol extract of the leaves. The structures of the new and known compounds were established by analyses of 1D- and 2D-NMR and mass spectrometry data.     

Morphology of female sex pheromone gland in the horse chestnut leafminer Cameraria ohridella (Lep., Gracillariidae)

Abstract: The internal and external morphology of the female sex pheromone gland in Cameraria ohridella Deschka & Dimic, an European pest on Aesculus hippocastanum L., has been investigated by histological and electron microscopic techniques. The gland consists of a single layer of modified epidermal cells in the dorsal part of the intersegmental membrane between the eighth and ninth abdominal segments and laterally extends to the posterior apophyses. The epithelium contains large columnar- and cone-shaped cells with basally situated nuclei. The cuticle, which covers the glandular region, has a wrapped appearance and is divided into a hyalin and thickened endocuticle and a thin outer epicuticle: it considerably expands when the gland is protruded and provides a sufficiently large surface for evaporation of the pheromone. The cuticle does not show any orifices of pore channels. In the retracted position, the gland is folded within the body cavity of the seventh and eighth abdominal segments but is exposed to the environment by extension of the abdominal tip along with female calling. In virgin females, pheromone glands are well developed at least within the first days after eclosion; if copulation occurs, glandular epithelia degenerate soon. According to the current classification, the glandular type of C. ohridella most easily is consistent with eversible dorsal scent folds that are widely distributed amongst diverse taxa of Lepidoptera. However, this is the first report on the morphology of pheromone glands in the Gracillariidae.     

Scent gland apparatus in the Western conifer seed bug Leptoglossus occidentalis Heidemann (Heteroptera: Coreidae)

The coreid Leptoglossus occidentalis is a Nearctic bug responsible for severe seed losses to pine orchards. When disturbed, adults and nymphs emit a defensive secretion deemed an allomone. Here we describe the gross morphology of the scent gland apparatus and the related evaporatory structures in nymphs and adults of L. occidentalis, through light microscopy and scanning electron microscopy. Adults of both sexes possess a metathoracic scent gland complex (MTG) including a central orange‐yellow reservoir and a pair of white lateral glands, connected by ducts to the reservoir. The MTG belongs to the diastomian type, with two ostioles located on the metathorax associated with a microsculptured cuticular accumulation area, i.e. evaporatory area, which can prevent the spread of the secretion on to non‐evaporative cuticle and increase scent fluid evaporation. A high number of male‐specific sternal gland pores were observed. These pores and associated glands are likely the source of an attractant pheromone, which could be extremely useful in monitoring and combating this invasive pest. In nymphs, MTG is replaced by two dorsal abdominal scent glands (DAGs) located between the 4th and the 6th urotergites. DAGs are reddish cuticle‐lined sacs with gland cells forming the gland wall; the scent substances are released through two orifices lying on the mid‐dorsal abdominal line between urotergites IV–V and V–VI. Also in nymphs, peculiar cuticular evaporatory areas surround both orifices.     

Fine Structure of the Esophagus of Males of Sarisodera hydrophila (Heteroderoidea)

The fine structure of the esophagus, including procorpus, metacorpus, isthmus, gland lobe, and esophago-intestinal junction, is examined in males of Sarisodera hydrophila. A cuticle-lined lumen extends most of the length of the esophagus, broadens to form a pump chamber in the metacorpus, and posteriorly is continuous with junctional complexes among four esophago-intestinal cells. These four cells are partially enveloped by the gland lobe which basically consists of three gland cells, one dorsal and two subventral. Each gland cell has an anterior process which opens into the lumen of the esophagus through a cuticle-lined duct. The dorsal gland joins the lumen in the anterior portion of the procorpus, whereas ducts of the subventral glands terminate at the base of the metacorpus pump chamber. The subventral glands are predominant in the posterior portion of the gland lobe and are partially ensheathed by a narrow portion of the dorsal gland which extends to within 5 μm of the posterior terminus of the gland lobe. Contents of the dorsal gland include primarily electron dense granules, although rough endoplasmic reticulum (RER) is predominant posteriorly. Secretory granules within the subventral glands vary in morphology and are evenly distributed throughout the two ceils among other organelles, including RER and a large Golgi apparatus. Innervation of the esophagus includes nerve processes which originate from several perikaryons (cell bodies) located in the anterior portion of the gland lobe. The esophagus of males of S. hydrophila is compared with that of other Heteroderoidea, Heterodera glycines and Meloidogyne incognita.     

三种蚤生殖系统的细微结构:雄性外生殖器的发育

本文研究了缓慢细蚤Leptopsylla segnis(Schonherr),不等单蚤Monopsyllus anisus(Rothschild)和猫栉首蚤指名亚种Ctenocephalides felis felis(Bouche)雄性外生殖器的结构,观察了从幼虫、前蛹、蛹至成虫各发育时期的雄性外生殖器的内部结构变化.对有争议的雄蚤上抱器的起源,雄蚤生殖孔的位置,雄性外生殖器芽内陷的腹节以及射精管横切面的细胞数目和阳茎背、腹杆的结构等问题进行了详细的观察和探讨.     

Functional anatomy of scent glands in Paranemastoma quadripunctatum (Opiliones,Dyspnoi, Nemastomatidae)

The morphological organization and functional anatomy of prosomal defensive (scent) glands in Paranemastoma quadripunctatum, a representative of the dyspnoid harvestmen, was investigated by means of histological semithin sections, software‐based 3D‐reconstruction and scanning electron microscopy. Scent glands comprise large, hollow sacs on either side of the prosoma, each of these opening to the outside via one orifice (ozopore) immediately above coxa I. In contrast to the situation known from laniatorean, cyphophthalmid and some eupnoid Opiliones, ozopores are not exposed but hidden in a depression (atrium), formed by a dorsal integumental fold of the carapace and the dorsal parts of coxae I. Glandular sacs are connected to ozopores via a short duct which is equipped with a specific closing mechanism in its distal part: A layer of modified epidermal cells forms a kind of pad‐like tissue, surrounding the duct like a valve. Several muscles attached to the anterior parts of the glandular reservoir and to the epithelial pad may be associated with ozopore‐opening. The actual mechanism of secretion discharge seems to be highly unusual and may be hypothesized on the basis of corroborating data from behavioral observations, scent gland anatomy and secretion chemistry as follows: Enteric fluid is considered to be directed towards the ozopores via cuticular grooves in the surface of the coxapophyses of legs I. Then, the fluid is sucked into the anterior part of the scent gland reservoirs by the action of dorsal dilator muscles that widen the reservoir and produce a short‐term negative pressure. After dilution/solution of the naphthoquinone‐rich scent gland contents, a secretion‐loaded fluid is thought to be discharged with the help of transversal compressor muscles. This is the first detailed study on the functional anatomy of scent glands and the mechanisms of secretion discharge in the Dyspnoi. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Two New Species of Afronurus (Ephemeroptera: Heptageniidae) from Vietnam

ABSTRACT Two new species of the heptageniid mayfly genus Afronurus Lestage, Afronurus meo sp. n. and Afronurus mnong sp. n., are described from Vietnam with line-drawings of key characters. The larvae of A. meo can be characterized by rowed minute spines in the posterior margin of the abdominal terga 1-9 and distinct markings on anterior head, dorsal femora, and abdominal terga. The larvae of A. mnong can be characterized by scattered simple stout setae on the anterior margin and dorsal surface of femora, rowed acute spines on the posterior margin of abdominal terga 1-9, and plain brown body color. Their diagnoses, material data, distributions, and habitat and biological data are provided.     

Regional Differentiation of Cortical Structures and Their Reorganization during Cell Division in Paramecium trichium

We observed the cell surface of Paramecium trichium using three different methods. A non-dividing paramecium's cell surface consisted of three major regions outside of the oral apparatus: a) an oral groove region, with 2-cilia-2-basal-body (2C-2BB) units; b) a posterior region, occupying 1/4 to 1/5 of the cell surface, with 1-cilium-l-basal-body (1C-1BB) units; c) the remainder, with l-cilium-2-basal-body (1C-2BB) units. Five kinds of region-specific cortical reorganization occurred prior to cytokinesis: the 2C-2BB and 1C-1BB units were not duplicated, while the 1C-2BB units were reorganized to 2C-2BB, 1C-2BB or 1C-1BB units. These reorganizations of the cell surface progressed from the fission line to the anterior in the prospective anterior daughter cell, and to the posterior in the prospective posterior daughter cell, and bilaterally from the old and also newly developing oral apparatus in both daughter cells. In contrast, the development of cilia and their associated structures in each of the cortical units always progressed from posterior to anterior. The present work also showed that two fission lines began to develop bilaterally from the oral primordium, and then they joined to become a single fission line at the dorsal surface.     

Cytotoxic lupane-type triterpenoids from Acacia mellifera

One new and eight previously described lupane-type metabolites were isolated for the first time from Acacia mellifera (Leguminosae). Based on spectral analyses, the structure of the new compound was elucidated as 28-hydroxy-3-oxo-lup-20-(29)-en-30-al (1), while the known compounds were identified as 3-oxo-lup-20-(29)-en-30-al (2), 3-hydroxy-lup-20-(29)-en-30-al (3), 28-hydroxy-lup-20-(29)-en-3-one (4), lupenone (5), lupeol (6), betulin (7), betulinic acid (8), and betulonic acid (9). Metabolites 2, 3, and 4 are reported for the first time in the Leguminosae family. The cytotoxicity of the isolated metabolites was evaluated on the NSCLC-N6 cell line, derived from a human non-small-cell bronchopulmonary carcinoma. Compounds 1 and 3 exhibited significant levels of activity.