Ultrastructure of the chemosensitive basiconic single-walled wall-pore sensilla on the antennae in adults and embryonic stages of Locusta migratoria L. (Insecta,Orthoptera)

Summary The structure and embryonic development of the two types (A, B) of basiconic sensilla on the antennae of Locusta migratoria were studied in material that had been cryofixed and freeze-substituted, or chemically fixed and dehydrated. Both types are single-walled wall-pore sensilla. Type-A sensilla comprise 20–30 sensory and 7 enveloping cells. One enveloping cell (thecogen cell secretes the dendrite sheath); four are trichogen cells, projections of which form the trichogen process during the 2nd embryonic molt. The trichogen cells form two concentric pairs proximally. Two tormogen cells secrete the cuticular socket of the sensillum. The dendritic outer segments of the sensory cells are branched. Bifurcate type-A sensilla have also been observed. Type-B sensilla comprise three sensory and four enveloping cells (one thecogen, two trichogen and one tormogen). The trichogen process is formed by the two trichogen cells, each of which gives rise to two projections. The trichogen cells are concentrically arranged. The dendritic outer segments of the sensory cells are unbranched. In the fully developed sensillum, all trichogen and tormogen cells border on the outer receptor lymph cavity. It is suggested that the multicellular organization of the type-A sensilla can be regarded as being advanced rather than primitive.Supported by the Dcutschc Forschungsgemeinschaft (SFB 4/G1)     

Ultrastructure and receptor cell responses of the antennal grooved peg sensilla of Triatoma infestans (Hemiptera: Reduviidae)

Ultrastructural examination of grooved-peg (GP) sensilla on the antenna of fifth instar Triatoma infestans nymphs by scanning electron microscopy and transmission electron microscopy reveal that they are 8–18 μm long with a diameter of about 2–2.8 μm at the non-articulated base. Some pegs have a terminal pore. These double-walled wall-pore (dw-wp) sensilla have an outer cuticular wall with 13–18 longitudinal grooves at the distal part of the peg. Groove channels are present at the bottom of the grooves from which radial spoke channels lead into the inner sensillum-lymph cavity. A dendrite sheath connects the tip of the thecogen cell to the inner cuticular wall thus forming separated outer and inner sensillum-lymph cavities. Four or five bipolar receptor cells are ensheathed successively within the GP sensilla by the thecogen cell, trichogen and tormogen cells. The inner dendritic segments of each sensory cell give rise at the ciliary constriction to an unbranched outer dendritic segment which can reach the tip of the sensillum.Electrophysiological recordings from the GP sensilla indicate that they house NH₃, short-chain carboxylic acid and short-chain aliphatic amine receptor cells and can be divided into three functional sub-types (GP 1–3). All GP sensilla carry a receptor cell excited by aliphatic amines, such as isobutylamine, a compound associated with vertebrate odour. GP type 1 and 2 sensilla house, in addition, an NH₃-excited cell whereas the type 2 sensilla also contains a short-chain carboxylic acid receptor. No cell particularly sensitive to either NH₃ or carboxylic acids was found in the grooved-peg type 3 sensilla. GP types 1, 2 and 3 represent ca. 36, 10 and 43% of the GP sensilla, respectively, whereas the remaining 11% contain receptor cells that manifest normal spontaneous activity but do not respond to any of the afore mentioned stimuli.     

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.     

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)     

Antennal thermo- and hygrosensitive sensilla in Antheraea pernyi (Lepidoptera,Saturniidae): ultrastructure and immunohistochemical localization of Na+,K+-ATPase

Summary In an attempt to identify and localize the components of voltage sources involved in sensory transduction in insect sensilla, the thermo-/hygrosensitive sensilla of the moth Antheraea pernyi were probed with a polyclonal antiserum against Na⁺,K⁺-ATPase in cryofixed and freeze-substituted preparations. The antiserum recognized epitopes on the cytoplasmic membranes of the dendritic inner segments and somata of the sensory cells and also on the cytoplasmic membranes of glial cells surrounding the initial axon segments. The findings support the current concept that ion pumps in the cytoplasmic membranes of the dendritic inner segments and somata of the sensory cells contribute to the maintenance of the resting potential of the sensory cells and to the driving forces generating the receptor currents in response to stimulation of the sensillum. Morphological features and immunohistochemical characteristics of the region of the initial axon segment are also discussed with respect to the initiation of action potentials in these sensilla.     

Structural and functional properties of the mechanoreceptors and chemoreceptors in the anterior oesophageal sensilla of the crayfish,Astacus astacus

Summary The anterior oesophageal sensilla (AOS) of Astacus astacus are short cuticular cylinders with a terminal pore. Three sensory cells belong to each AOS, two of them having a dendritic ciliary segment of about 2 m length (A-cells), the third having this segment about 12 m long (B-cell). The ciliary A-tubules in the B-cells only possess arms and an electron-dense core. The dendritic inner segments of the A-cells terminate 3–6 m distal of the B-cells. The dendritic tips of most A-cells are connected by desmosomes. All dendritic inner segments contain a ciliary rootlet and are accompanied by a scolopale within the innermost enveloping cell. There are four enveloping cells, three of which form thin subepithelial columns together with the enclosed sensory cells. Recordings from the posterior branch of the anterior oesophageal nerve containing the axons of the AOS revealed the presence of three types of sensory cells, two being chemosensitive and one mechanosensitive. One chemoreceptor is specifically sensitive to nicotinamide, but responded also to -NAD, 6-aminonicotinamide, nicotinamide methyl esther and nicotin. It was blocked by p(4)-acetylpyridine. The second chemoreceptor responded only to crayfish gastric fluid. The mechanoreceptors reacted to stretch of the oesophageal wall adapting only slowly to maintained stimuli. It is assumed that the A-cells are the chemosensitive cells and the B-cells the mechanosensitive ones. The latter show only a small number of modality-specific characteristics. Several structural features appear to be correlated with the location of the AOS within a flexible surface, which undergoes considerable dilation.Supported by the Deutsche Forschungsgemeinschaft (H.A.:SFB 4/G1; H.H.: HA 1201/1-4)     

Differentiation of the thermo-/hygrosensitive (no-pore) sensilla on the antenna of Antheraea pernyi (Lepidoptera,Saturniidae): a study of cryofixed material

Summary The morphogenesis of the thermo- and hygro-sensitive sensilla styloconica of Antheraea pernyi was studied, exclusively by cryomethods, during the second half of pupal development. The three major processes taking place during this period are (1) the differentiation of the dendritic outer segments of the sensory cells, especially of the lamellated type-2 receptor, (2) the formation of the receptor-lymph cavities, (3) the formation of tubular structures of unknown function in the inner receptor-lymph cavity, and (4) the elongation of the dendrite sheath. The formation of lamellae in the type-2 dendritic outer segment is achieved by the enfolding of its originally cylindrical cytoplasmic membrane. Autocellular junctions, previously described in the sensilla of adult animals, are found to join the forming lamellae. Close similarities between the junctions and smooth septate junctions are demonstrated. Both the extensive inner and outer receptor-lymph cavities are formed by invagination and folding of the apical cytoplasmic membranes of the three enveloping cells. Formation starts at the most apical projection of the cells and proceeds in a proximal direction. Up to 4-m-long tubular structures appear, exclusively in developmental stages, in the inner receptor-lymph cavity. They are composed of plasma membranes whose inner surface is studded with regularly spaced electron-dense particles. Contacts with the cytoplasmic membrane of the innermost enveloping cell demonstrate that the structures are composed of lipid membranes. During elongation of the dendrite sheath, which in these sensilla is apically attached to the hair wall, an 2-m-long growth-zone is observed at its proximal end. By addition of sheath-forming material to the growth-zone, the latter continuously moves proximally until the sheath is completed.     

浙江省大雷山刺齿虫兆属一新种描述（弹尾纲：长角虫兆科）

记述浙江省大雷山刺齿虫兆属 1 新种,张氏刺齿虫兆Homidia zhangi sp. nov., 该种鉴别特征有体色,光滑下唇毛 L1,腹部第 4 节后侧中间 3+3 根大刚毛及长度相近的特化毛,腹部第 1 节和第 5 节特化毛的相对位置以及腹管侧瓣和弹器端区的大量纤毛状刚毛。模式标本保存于南京大学生命科学学院和台州学院生命科学学院。     

Hygro- and thermoreceptors in tip-pore sensilla of the tarsal organ of the spider Cupiennius salei: innervation and central projection

The hygro- and thermoreceptive tarsal organ in the wandering spider Cupiennius salei is located on the tarsus of each walking leg and pedipalp, and consists of a tiny air-filled capsule in the cuticle. This capsule communicates with the outside world through a small aperture and contains seven nipple-shaped sensilla, each with a pore at its tip. In both their external morphology and internal structure, the sensilla are indistinguishable, although one sensillum is innervated by only two sensory cells, whereas the other six sensilla contain three sensory cells. Their dendrites are unbranched and terminate at the tip-pore, where they are enveloped by amorphous material that appears to limit their exposure to the atmosphere. Cobalt fillings reveal that each tarsal organ projects to three different areas within the suboesophageal ganglionic mass: (1) the sensory longitudinal tract 3 and 4; (2) the corresponding pedipalpal or leg ganglion; (3) a structured neuropil (here termed the Blumenthal neuropil) beneath the oesophagus. The multiple representation of sensory afferents from each tarsal organ in different regions of the suboesophageal ganglionic mass suggests parallel processing of hygro-/thermoreceptive information.     

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)     

Metamorphic changes in the central projects of hair sensilla inTenebrio molitor L. (Insecta: Coleoptera)

Summary This paper describes the afferent projections of hair sensilla of the pro- and mesothoracic legs and the lateral thoracic sclerites of larval and adultTenebrio molitor and the corresponding set of pupal hair sensilla. The sensory neurons that innervate the hair sensilla of larval or adult insects project somatotopically into the thoracic neuropil. Different types of sensilla on the same region of the body surface project to the same zone of the ipsilateral thoracic ventral neuropil but exhibit different arborization patterns. Although there is a profound reorganization of body surface sensilla, the basic somatotopic layout of the larva is maintained in the adult. The sensory neurons that innervate the pupal hair sensilla possess central projections similar to those of the corresponding adult sensory neurons. The central projections of pupal sensory neurons are somatotopically oriented. Their projection pattern is serially homologous in the thoracic and the abdominal ganglia. The central projection pattern of the described pupal sensory neurons is constant throughout pupation. MAb 22C10 immunoreactivity allows an estimate of the timing of the early differentiation of the imaginal sensory neurons originating during pupation. Ablation experiments indicate that pupal sensory neurons influence the central projection pattern of the differentiating imaginal sensory neurons.     

中国东北地区滨棘虫兆族一新种及一新记录种（弹尾纲：棘虫兆科）

记述采自中国东北地区棘虫兆科滨棘虫兆族Thalassaphorurini1新种:宋氏异棘虫兆Allonyc hiurus songi sp.nov.及1新记录种:利富滨棘虫兆Thalassaphorura lifouensis(Thibaud&Weiner,1997),附详细描述及特征图。研究标本保存于中国科学院东北地理与农业生态研究所湿地生态与环境重点实验室。     

The formation of intercellular junctions in insect stem cell progeny (cockroach intestinal epithelium)

Summary The stages in the development of intercellular junctions have been followed in the mesenteric caecal cells of the cockroach midgut, where two types of mature cell, the columnar and the secretory, exist. Nests of undifferentiated replacement cells occur at intervals along the basal lamina, consisting of central, dividing cells and peripheral semi-lunar cells; the former act as proliferative stem cells to give rise to either pre-columnar or pre-secretory cells. The semi-lunar cells are pre-columnar and produce an attenuated process which gradually projects up to the luminal surface, producing microvilli and a dense extracellular substance en route. Intercellular gap junctions appear between these maturing columnar cell borders first, while septate junctions differentiate later; these are assembled from two different sets of intramembranous particle which become organized into either plaques or rows in parallel alignment, possibly mediated by actin filaments and microtubules. The pre-secretory cells, which are much fewer in number, remain associated only with the basal lamina and never reach the lumen; they develop into one of three distinct mature secretory cell types which release their secretory product in different ways. Offprint requests to: N.J. Lane     

Ontogeny of the antennal glands in the crayfish Astacus leptodactylus (Crustacea, Decapoda): anatomical and cell differentiation

The ontogeny of the antennal glands was studied during the embryonic and post-embryonic development of Astacus leptodactylus. The future glands arising from undifferentiated columnar cells were detectable at the metanauplius stage EI 150 m (EI: eye index; approximately 440 m at hatching). The tubule and labyrinth differentiated in embryos at EI 190 m, and the bladder and coelomosac at EI 250 m. At EI 350 m, the tubule lengthened and divided into proximal and distal sub-regions. In later stages, the gland retained the same morpho-anatomy but the differentiation and size of each part increased. The cells of the coelomosac displayed the cytological features of podocytes in late embryonic development at EI 440 m. Only small apical microvilli and a few mitochondria were observable in the labyrinth cells at EI 250 m; by EI 440 m, these cells presented well-shaped apical microvilli, formed bodies, basal infoldings and mitochondria. In the cells of the tubules and bladder, mitochondria and basal infoldings occurred at EI 440 m and EI 250 m, respectively. The differentiation of the tubules and bladder cells suggested that they were involved in active transport at EI 440 m. Following hatching, the differentiation of the cells and the size of the glands increased. The ontogeny of the antennal glands thus starts in early embryos, the specific cellular functional features being differentiated in the various parts of the glands by EI 440 m. The antennal glands are probably functional just before hatching, i.e., before the juveniles are confronted with the low osmolality of freshwater.Thanks are due to the University of Tarbiat Modarres and Ministry of Science, Research and Technology, Islamic Republic of Iran, for financial aid and support. Special thanks are also extended to the Société Française dExportation des Ressources Educatives (SFERE) for a scholarship to S.K.     

Localisation of pancreatic polypeptide (PP)-like immunoreactive material in neurones of the brain of the blowfly,Calliphora erythrocephala (Diptera)

Summary The brain of the blowfly, Calliphora erythrocephala, has been studied by means of the peroxidase-antiperoxidase immunocytochemical method, with the use of antibodies to bovine pancreatic polypeptide (BPP). A number of immunoreactive neurones have been localised, some corresponding to neurones previously identified tentatively as neurosecretory. This finding is further evidence that biologically active peptides, previously considered to be vertebrate, also exist in invertebrates. It also supports the concept of their evolutionary origin in nervous tissue.     

Fine structure of a sensory organ in the arista of Drosophila melanogaster and some other dipterans

Summary The arista, a characteristic appendage of dipteran antennae, consists of 2 short segments at the base and a long distal shaft. A small sensory ganglion, from which arises the aristal nerve, is located proximally in the shaft. The fine structure of the aristal sensory organ was studied in detail in the fruitfly (Drosophila) and for comparison in the housefly (Musca) and the blowfly (Calliphora). In Drosophila, the aristal sense organ consists of 3 identical sensilla that terminate in the hemolymph space of the aristal shaft, and not in an external cuticular apparatus. Each sensillum comprises 2 bipolar neurons and 2 sheath cells; a third sheath cell envelops the somata of all six neurons of the ganglion. The neurons have long slender dendrites with the usual subdivision into an inner and an outer segment. One of the outer segments is highly lamellated and bears small particles (BOSS-structures) on the outside of its cell membrane; the other outer segment is unbranched and has a small diameter. The fine structure of the first dendrite is strongly reminiscent of thermoreceptors known from the antennae of other insects. These thermoreceptors are often coupled with hygroreceptors; however, we can only speculate whether the second dendrite of the aristal organ also has this function. Our present results argue against mechanoreceptive functions, as formerly postulated. The aristal sense organs in Musca and Calliphora are similar to those in Drosophila, but contain more sensilla (12 in Musca, 18 in Calliphora).     

The antennae of neopseustid moths: Morphology and phylogenetic implications, with special reference to the sensilla (Insecta, Lepidoptera, Neopseustidae)

The antennae of Lepidoptera Neopseustidae were examined with the scanning electron microscope. The studied species, Nematocentropus cfr. omeiensis, Neopseustis meyricki, Synempora andesae, Apoplania valdiviana and Apoplania penai possess nine types of antennal flagellum sensilla: multiporous large sensilla basiconica, multiporous thin sensilla basiconica, multiporous small sensilla basiconica, multiporous sensilla trichodea, multiporous sensilla coeloconica; uniporous sensilla chaetica; aporous sensilla chaetica, aporous stylus-shaped sensilla chaetica, aporous sensilla styloconica.The multiporous sensillum type here termed “multiporous large sensillum basiconicum” is unknown from other Lepidoptera and probably constitutes an autapomorphy of the family Neopseustidae. This sensillum type is remarkable by having a single base in female Apoplania and Synempora while in male Apoplania it has a bifid or trifid base, and in male Synempora it is composed of two or three incompletely separated hairs. This may be the first recorded example of a sexually dimorphic lepidopteran sensillum type. The stylus-shaped sensillum chaeticum is a primitive type which occurs only in some lower Lepidoptera.     

The fine structure of thermo-/hygrosensitive sensilla in the silkmoth Bombyx mori: Receptor membrane substructure and sensory cell contacts

Summary The thermo-/hygroreceptive sensilla styloconica of the silkmoth Bombyx mori are located on the tips of the antennal branches. A small poreless cuticular peg is innervated by three sensory cells. The outer dendritic segments of two type-1 receptor cells, the presumed hygroreceptors, almost completely fill the peg lumen and are in close contact with each other. The outer dendritic segment of the third (type-2) receptor cell, the presumed thermoreceptor, forms lamellae below the peg base. The membranes of these lamellae are studded with knobs in orthogonal array, protruding into the extracellular space with the same orientation on facing lamellae. This Bossy Orthogonal Surface Substructure (BOSS) is assumed to play a role in thermoreceptor function. Contacts are observed between the outer dendritic segments, between the inner dendritic segments immediately below the ciliary segments, and between the sensory cell somata. These contacts, which are not found in the olfactory sensilla (s. trichodea and basiconica) of this species, indicate electrical interactions between the three sensory cells of the styloconic sensillum and possibly are involved in the antagonistic and/or bimodal response characteristics of thermo-/hygroreceptor cells.     

Morphology of neurones associated with the antennal heart of Periplaneta americana (Blattodea,Insecta)

Summary Innervation of the antennal heart, an independent accessory circulatory motor in the head of insects, was investigated in the cockroach Periplaneta americana by use of axonal cobalt filling and transmission electron microscopy. The muscles associated with this organ are innervated by neurones located in a part of the suboesophageal ganglion, generally considered to be formed by the mandibular neuromere. Dorsal unpaired median (DUM) and paired contralateral neurones were stained. The axons of all these neurones run along the circumoesophageal connectives and through the paired nervus corporis cardiaci III into the corpora cardiaca. They pass through these organs forming fine arborizations there and exit anteriorly as a small pair of nerves which terminate at the antennal heart-dilator muscles. Numerous branches of these nerves extend beyond the lateral borders of the large transverse dilator muscle and terminate in the ampullar walls of the antennal heart. These neurosecretory fibres form neurohaemal areas which obviously release their products into the haemolymph, which is pumped into the antennae. The possible functions of the neurones associated with the antennal heart are discussed with respect to both, their role as a modulatory input for the circulatory motor and as a neurohormonal release site.