The chemosensory behaviour of Lithobius forficatus. 1. Evidence for a pheromone released by the coxal organs (Myriapoda: Chilopoda)

The behaviour of the Common brown centipede, Lithobius forficatus (.L.) is investigated with reference to the function of the coxal organs which are a peculiar and characteristic feature of Chilopoda. Previous studies of these organs in centipedes have been largely morphological and have relied on ultrastructural comparisons with other arthropod epithelia of disparate functions. Evidence is presented which suggests that the coxal pores release a sex-specific pheromone. The nature of intraspecific interactions in L. forficatus is discussed.     

The chemosensory behaviour of Lithobius forficatus (Myriapoda: Chilopoda). 2. Bioassay and chemistry of the coxal pheromone

The common brown centipede, Lithobius jorJcatus (L.), releases a sex-specific pheromone which emanates from distinctive cuticular openings on the coxae of the hind legs-the coxal pores (Littlewood & Blower, 1987). The pheromone can be isolated from tissue extracts of the coxal organs (surrounding the coxal pores) and from substrata which have been 'marked by male or female L. forjcatus and presented to centipedes in the form of a behavioural bio-assay.
In this paper, further evidence is presented for a centipede pheromone, the chemical components of male and female secretions are separated using Gas Layer Chromatography (GLC), a qualitative chemical analysis is described and the nature of the pheromone is discussed.     

Chilopod coxal organs: morphological considerations with reference to function

A comparative account of the morphology of chilopod coxal organs is given with special reference to their possible function.
Allometric relationships which exist between the size of the centipede and the number of coxal pores and organs are discussed in the context of size-dependent physiological constraints which may affect the function of the coxal organs.     

Ultrastructural organization of the anal organs in the anal capsule of Craterostigmus tasmanianus Pocock, 1902 (Chilopoda, Craterostigmomorpha)

We describe the ultrastructural organization of the anal organs of Craterostigmus tasmanianus, which are located on the ventral side of the bivalvular anal capsule. Each part of the capsule bears four pore fields with several anal pores. The pores lead into a pore canal, which is surrounded by the single-layered epithelium of the anal organs. Each anal organ is composed of four different cell types: transporting cells of the main epithelium, junctional cells, isolated epidermal glands, and the cells forming the pore canal. The transporting cells exhibit infoldings of the outer cell membranes, forming a basal labyrinth and a poorly developed apical complex. The cells are covered by a specialized cuticle with a widened subcuticular layer. Only the cuticle of the main epithelium is covered by a mucous layer, secreted by the epidermal glands. The ultrastructural organization of the anal organ is comparable to the coxal and anal organs of other pleurostigmophoran Chilopoda. It is likely that the coxal and anal organs of the Pleurostigmophora are homologous, due to their identical ultrastructural organization. Differences concerning the location on the trunk of Pleurostigmophora are not sufficient to reject a hypothesis of homology. Anal organs are found not only in Craterostigmomorpha, but also in most adult Geophilomorpha, and in larvae and most adults of Lithobiomorpha. The anal organs of C. tasmanianus are thought to play an important role in the uptake of atmospheric water. J. Morphol.     

Coxal organs in geophilomorpha (Chilopoda). Organization and fine structure of the transporting epithelium

Summary The coxal organs of different Geophilomorpha were studied by scanning and by transmission electron microscopy.1) The coxae of the last trunk-segment contain pores in different arrangements and numbers. They are the openings of the coxal organs.2) The coxal organs are formed by four different cell types: the main epithelium consists of radially arranged transporting cells, surrounded by junctional cells, gland cells, and the cells of the pore channel.3) The cells of the transporting epithelium show an enlargement of the apical and basal surface. Deep and narrow extracellular channels of the apical infoldings are closely associated by mitochondria (plasmalemma-mitochondrial complexes). The epithelium is covered by a prominent cuticle with a spacious subcuticle.4) A distinct mucous layer covers the cuticle of the transporting epithelia, and is secreted by the gland cells.5) A small cellular sheath separates the epithelium of the coxal organ against the haemolymph.6) The possible function of the coxal organs in ion and fluid transport is discussed.     

Seasonal fluctuations in the protozoan parasites of the centipedes Lithobius variegatusand Lithobius forficatus in a Yorkshire woodland

The seasonal variation in percentage parasitization and in the average number of individuals of Echinomera spp. in Lithobius forficatus and L. variegatus is described and discussed. It is suggested that infection by Echinomera hispida may account for the low numbers of L.forficatus in damp woodlands.
Coccidia occur only in L. variegatus and the incidence of metazoan parasites is very low.     

THE FOOD AND REPRODUCTIVE CYCLES OF THE CENTIPEDES LITHOBIUS VARIEGATUS AND LITHOBIUS FORFICATUS IN A YORKSHIRE WOODLAND

Lithobius variegatus and L. forficatus occupy similar ecological niches and frequently occur together although there are differences in their distribution in the British Isles. An investigation of their food and reproductive cycles, being a preliminary step towards a further understanding of their ecological relationships, is described.
Lithobiomorph centipedes have hitherto been regarded as wholly carnivorous, but it is shown that these species feed on litter as well as small litter animals, the presence of litter in the guts of these species not being connected with the presence of animal remains. Whereas L. forficatus takes litter throughout the year, L. variegatus does so mainly in the winter.
Both species appear to lay eggs for a considerable part of the year, though there appears to be only a short period of sperm transfer in spring. It is suggested that the long oviposition period may prevent the loss of the entire brood in a dry summer. The succession of epimorph stadia in L. variegatus is described.
L. forficatus was more common than L. variegatus in the drior part of the woods studied.     

Comparative ultrastructure of coxal glands in unfed larvae of Leptotrombidium orientale (Schluger, 1948) (Trombiculidae) and Hydryphantes ruber (de Geer, 1778) (Hydryphantidae)

Coxal glands of unfed larvae Leptotrombidium orientale (Schluger, 1948) (Trombiculidae), a terrestrial mite parasitizing vertebrates, and Hydryphantes ruber (de Geer, 1778) (Hydryphantidae), a water mite parasitizing insects were studied using transmission electron microscopy. In both species, the coxal glands are represented by a paired tubular organ extending on the sides of the brain from the mouthparts to the frontal midgut wall and are formed of the cells arranged around the central lumen. As in other Parasitengona, the coxal glands are devoid of a proximal sacculus. The excretory duct, joining with ducts of the prosomal salivary glands constitutes the common podocephalic duct, opening into the subcheliceral space. The coxal glands of L. orientale are composed of a distal tubule with a basal labyrinth, an intermediate segment without labyrinth, and a proximal tubule bearing tight microvilli on the apical cell surface and coiled around the intermediate segment. The coxal glands of H. ruber mainly consist of the uniformly organized proximal tubule with apical microvilli of the cells lacking the basal labyrinth. This tubule shows several loops running backward and forward in a vertical plane on the side of the brain. In contrast to L. orientale , larvae of H. ruber reveal a terminal cuticular sac/bladder for accumulation of secreted fluids. Organization of the coxal glands depends on the ecological conditions of mites. Larvae of terrestrial L. orientale possess distal tubule functioning in re‐absorption of ions and water. Conversely, water mite larvae H. ruber need to evacuate of the water excess, so the filtrating proximal tubule is prominent.     

The contributions of diverse sense organs to the control of leg movement by a walking insect

Summary During locomotion, stick insectsCarausius morosus, place the tarsus of the rear leg near the tarsus of the ipsilateral middle leg, whatever the position of the latter. This adjustment by the hind leg requires that it receive information on the actual position of the middle leg tarsus. It is shown by ablation experiments that such information is contributed by the following proprioceptors of the middle leg: the ventral and dorsal coxal hairplates, the coxal hair rows, the trochanteral hairplate and the femoral chordotonal organ. Additional information comes from other, as yet unidentified, sense organs. Several alternatives are considered to explain how the signals from the diverse sense organs of the subcoxal joint might be combined in computing the target position for the protracting hind leg. The experimental results support the hypothesis that the signals are added nonlinearly and that a signal deviating from the majority pattern is weighted less.Abbreviations cxHPu ventral coxal hairplate - cxHPd dorsal coxal hairplate - trHP trochanteral hairplate - HR hair row - feCO femoral chordotonal organ - AEP anterior extreme position     

The coxal glands of geophilomorpha (chilopoda): Organs of osmoregulation

Summary The organs terminating at the coxal pores of the tug-legs of Geophilomorpha are not repugnatorial glands, but possess typical transport epithelia with deep apical and basal infoldings of the cell membranes, between which numerous large mitochondria are located. Many transport vesicles are found in the basal region but fewer in the apical cytoplasm. The apex is characterized by bundles of longitudinally oriented microtubules, sparse endoplasmic reticulum and free ribosomes. Single neurosecretory axons with synaptoid areas are scattered among the cells. It is suggested that the coxal organs have a diuretic function in moist habitats and an antidiuretic effect in arid environments. The switch-over is evidently controlled by a neuroendocrine mechanism.     

Comparative morphology,functions, and homologies of the coxal glands in oribatid mites (Arachnida: Acari)

1. Forty-eight species of oribatids in 37 families representing most of the superfamilies were collected from various environments (littoral, salt marsh, litter, sod, and freshwater) and sectioned. 2. The coxal gland is composed of a sacculus and a labyrinth in all stages of all oribatid species. Muscles, originating on the body wall, insert at several points on the thin-walled sacculus which opens into the labyrinth. The labyrinth has an internal, chitinous supporting skeleton. The type A labyrinth has 3–180° bends, producing four parallel regions, and occurs in all inferior oribatids. The type B labyrinth has 1–180° bend, producing two parallel regions, and occurs in all superior oribatids. The coxal gland duct and the lateral gland duct join, penetrate the body wall, and empty into the posterior end of the podocephalic canal. All oribatids have lateral accessory glands, but only inferior oribatids have rostral and medial glands. Three ductless coxendral bodies are always present. 3. The labyrinth length in oribatids is correlated with body size and the environment of the species. Oribatids from sod, leaflitter, or moss show a simple correlation of labyrinth length (X) to total body length (Y) where Y = 4.64X. Freshwater species have a labyrinth length greater than that of comparably sized terrestrial species and salt water (littoral) species have a labyrinth length less than that of comparably sized terrestrial species. There is a greater reduction in labyrinth length in species restricted to salt marshes than in species not restricted to salt marshes. 4. The probable function of oribatid coxal glands is osmoregulation. Hemolymph filtration would occur across the sacculus by positive hemolymph pressure and contraction of the sacculus muscles. Resorption of ions would occur in the labyrinth, which is noncollapsible due to the internal skeleton. The hypothesis is that in freshwater species the rate of filtration is high and resorption of ions would have to be very efficient, therefore they have an elongated labyrinth; but in salt water species water loss must be minimized and preservation of ions would be a disadvantage, therefore they have a shortened labyrinth. Excre ion may also be a function of the coxal glands. The lateral gland may possibly function as an endocrine gland involved with production of a molting hormone. The rostral glands in inferior oribatids may have a salivary function. 5. The coxal glands of Peripatus, some millipedes, apterygote insects, decapod crustaceans, and all arachnid orders are homologous. The Tetrastigmata, Notostigmata, Cryptostigmata, and soft ticks have typical arachnid coxal glands. The coxal glands of higher Prostigmata may be modified into salivary, silk, or venom glands. The coxal glands in Mesostigmata, Astigmata, and hard ticks are lacking or highly modified.     

The ultrastructure of the sternal gills forming a striking contrast with the coxal gills in a fresh-water amphipod (Crustacea)

Sternomoera yezoensis has specialized sterna with 21 sternal gills in addition to six pairs of coxal gills. Despite a common high permeability to chloride ions, the cpithelia of these two kinds of gills are diametrically opposed in the polarity of the cell membranemitochondria complex. The coxal gill epithelium (4-6 mum thick) is characterized by a welldeveloped AIS (apical infolding system) associated with a huge number of large mitochondria. The AIS exceeds two-thirds of the epithelial thickness and forms a highly sophisticated, subcuticular labyrinth. On the contrary, the sternal gill epithelium, an extension of the sternal epithelium proper, is extremely thick (10-15 mum) and is characterized by a very deep BIS (basolateral infolding system) associated with numerous slender mitochondria. The BIS reaches nine-tenths of the epithelial thickness and forms a giant, baso-lateral labyrinth. Shallower, less elaborate AIS and BIS without mitochondrial association originate from the opposite sides of these epithelia. Although AIS and BIS interpenetrate in the sternal gill epithelium, they never communicate. The results indicate that in addition to the coxal gills, the sterna with Ihe sternal gills function as transporting as well as respiratory organs, though the functional difference between these two kinds of gills remains to be elucidated.     

Coxal organs of Lithobius forficatus (Myriapoda,Chilopoda)

Summary In Lithobius forficatus each of the coxae of the four posterior trunk segments bear a pore field with several coxal pores. The surrounding single-layered epithelium is composed of four different cell types: the main epithelial cells having a fine-structural organization of transport cells with deep apical and basal folds of the cell surfaces and plasmalemma-mitochondrial complexes, junctional cells, exocrine glands, and the wall cells of the pore channel. The entire epithelium is separated from the hemolymph by an inner cellular sheath. It is assumed that the coxal organs participate in fluid uptake.     

Chemoreceptor sensillum structure in Limulus

The chemoreceptors of Limulus polyphemus (L.) are polyneuronal sensilla found in the spines of the coxal gnathobases of each walking leg, the spines of the chilarial appendages, and the chelae of all the limbs. Each sensillum contains 6–15 bipolar sensory cells that share a single pore in the cuticle. The dendrites of the sensory cells of each sensillum course to the cuticle together. These attenuate sharply and enter a canal in the cuticle as a very narrow terminal thread. The dendrites retain their identity in the thread, but with the light microscope, they are usually not visible individually. Each thread, consisting of 6–15 dendrites, is accompanied to the cuticular surface by a cuticular tubule found within the canal. The chemoreceptor sensilla of the gnathobase, chilarium, and chela, the temperature organs of Patten, and the flabellar receptor organs all have the same basic organization. In general this is the same structural plan shown by chemoreceptors of other arthropods. Several different mechanisms of peripheral physiological interaction among receptor cells are possible with a sensillum organization like that described here for Limulus.     

Transport-active organs within the ’ano-genital’ capsule of Craterostigmus tasmanianus (Chilopoda, Craterostigmomorpha)

 In Craterostigmus tasmanianus, first results of the cellular organization of anal organs within the ’ano-genital’ capsule are presented. Each valve of the ’ano-genital’ capsule bears four pore fields ventrally, each of them consisting of several pore openings of the anal organs. The pores lead into a cuticle-lined pore channel, the base of which is surrounded by a single-layered epithelium that is composed of three different cell types. The main epithelium consists of radially arranged transport-active cells surrounded by exocrine cells, and the cells of the pore channel. The cells of the transporting epithelium show deep invaginations of the apical and basal cell surfaces and plasmalemma-mitochondrial complexes. These cells are covered by a specialized cuticle with a prominent subcuticle. Exocrine glands secrete a mucous layer on the cuticle of the main epithelium. The type of anal organ present in Craterostigmus tasmanianus shows similarities to coxal and anal organs found in other Pleurostigmophora in the chilopods. The possible function of the anal organs in uptaking water vapour is discussed. It is appropriate to call the organs within the ’ano-genital’ capsule of Craterostigmus tasmanianus ”anal organs”, as components of the genital segments are not involved. Accepted: 17 November 1996     

The peripheral and central nervous organization of the locust coxo–trochanteral joint

The micromorphology of the locust coxo–trochanteral joint was examined in cobalt-stained material. Peripheral nervous system, musculature, and internal proprioceptors—two strand receptors and a muscle receptor organ—of the metathoracic coxa are compared with those of the pro- and mesothoracic legs. The number and position of trochanter levator and depressor motoneurons as well as the central projections of coxal sense organs are described. Evidence for a femoro–tibial strand receptor was obtained by tracing the path of a particular nerve branch.     

Biochemical changes in tissue composition of Periplaneta americana (Linn.) acclimated to high and low temperatures

1. 1.|Cold acclimation apparently favours an increase of water content in fat body, but not in coxal muscle, of cockroaches.

2. 2.|A remarkable enhancement in the accumulation of total protein in fat body characterizes the cold acclimation of cockroaches, particularly adult males (175% increase in protein/DNA ratio). The increase in protein content of coxal muscle during acclimination to 15°C, observed in nymphs (16%) and males (16%) but not in females, is less pronounced than that of fat body.

3. 3.|A diminution (28–32%) in the free amino acid/DNA ratio due to cold acclimation has been recorded in both coxal muscle and fat body of nymphs and females, but not in males.

4. 4.|No qualitative change occurs in the free amino acid spectrum of haemolymph and tissues of this insect during acclimation to 15 and 35°C.

5. 5.|An augmentation (15–30%) of the RNA/DNA ratio occurs in fat body and coxal muscle of nymphs and males but in fat body alone of females following cold acclimation.

6. 6.|The glycogen reserve has been shown to increase by up to 30% in fat body and coxal muscle of cold acclimated cockroaches compared to warm acclimated ones.

The proprioceptive function of a complex chordotonal organ associated with the mesothoracic coxa in locusts

Summary A system of chordotonal organs in the locust mesothorax consists of four subunits one of which connects to the coxa. Proprioceptive afferents from the scoloparia record the rotatory movements of the coxa. Mechanical stimulation of the sensory system by sinusoidal stretch or movements mimicking stretch as in natural walking of the locust elicits reflex activation of coxal motoneurones. Both assistance and resistance reflexes to imposed movements occur, but their intensity can vary from periods of suppression below firing threshold in a motoneurone to recruitment of additional motoneurones to the same muscle. It is concluded that some of these reflexes recorded in isolated preparations can also occur in freely walking animals where they should contribute to the muscular coordination of transitions between antagonistic movements.Abbreviations aCO, cCO, pCO, vCO anterior, coxal, posterior, ventral chordotonal organ - COS chordotonal organ system - pm-al postero-median to anterior-lateral     

Fine structure and function of the coxal glands of lithobiomorph centipedes: Lithobius forficatus and L. crassipes (Chilopoda,Lithobiidae)

The ultrastructure of the coxal glands and associated tissues in the centipedes Lithobius forficatus and Lithobius crassipes has been examined in the light of two contrasting functional hypotheses postulated by different authors. Lithobiomorph chilopods possess eight sets of pores on the posterioventral border of the coxal podomeres of leg pairs 12–15 in adult (maturus) and subadult (pseudomaturus) stadia. A modified cuticular hypodermis, known as the coxal gland, surrounds the distal portion of each blindended pore. Each gland is made up of cells which contain large numbers of hypertrophied mitochondria and a highly folded apical and basal plasma membrane. The similarity of the coxal gland to so called “transporting epithelia” is discussed and further comparisons are made between these and secretory glands in arthropods. A careful consideration of both functional hypotheses (osmoregulation or pheromone release) has revealed the possibility that the coxal gland may encompass both functions.