Internal and external morphology of the deutonymph of Caloglyphus boharti (Arachnida: Acari)

A band of flexible cuticle encircles the deutonymph, separating the dorsal and ventral plates. The coxae are large, flat and fused with one another to form most of the ventor. Individual coxal margins are redefined as sternites, epimerites or simply apodemes according to which margins fuse with which others. A given area of cuticle may have patches of dark or light cuticle not corresponding to particular structures or cuticular contours; this is a source of confusion to taxonomists. Each leg has a dicondylic coxal-trochantal (adduction-abduction) and trochantal-femoral (promotion-remotion) joint with opposing muscles. The three more distal monocondylic joints (flexion-extension) have only flexor muscles; extension is by increased haemolymph pressure. The five apodemes of the sucker plate provide rigidity; the four suckers attach by a flexible cuticular ring to a solid flange or socket in the sucker plate. The sucker muscles attach to the center of each sucker. The flat, external face of the sucker plate apodemes may complement sucker action by adhesion. Coxal discs and sucker plate discs are identical, contain birefringent cuticular elements, and are considered modified setae. Functional mouthparts and a pharynx are lacking, but a cheliceral anlage is present. The esophagus, midgut and caecae, and malpighian tubules are lumenless and the cells small. The hindgut has a lumen, larger cells and opens externally via the anus. Whereas the digestive tract is regressed, the reproductive system is yet incompletely developed. In older deutonymphs anlagen of ducts, accessory glands and gonads are discernible. The nature of the haemocoel and peritoneum remains nuclear. The central nerve mass is conspicuously large for the size of the deutonymph. The supraesophageal ganglion gives rise to the cheliceral nerves; all other nerves arise from the subesophageal ganglion. Most major nerves were traced to the effector organs. The muscles are divided into leg, dorso-ventral (derived from coxal muscles), dorsal, sucker, and anogenital muscles. The trochantal adductor muscles originate on an endosternite, which is supported by muscles running to the dorsal hysterosoma. The dorso-ventral and propodosomal retractor muscles affect haemolymph pressure. The massive sucker retractor muscles are unique to this instar. Anogenital muscles are not well developed.     

Morphology and ultrastructure of the esophagus during the ontogeny of the spider crab Maja brachydactyla (Decapoda,Brachyura, Majidae)

The esophagus of the eucrustaceans is known as a short tube that connects the mouth with the stomach but has generally received little attention by the carcinologists, especially during the larval stages. By this reason, the present study is focused on the morphology and ultrastructure of the esophagus in the brachyuran Maja brachydactyla during the larval development and adult stage. The esophagus shows internally four longitudinal folds. The simple columnar epithelium is covered by a thick cuticle. The epithelial cells of the adults are intensively interdigitated and show abundant apical mitochondria and bundles of filamentous structures. The cuticle surface has microspines and mutually exclusive pores. Three muscle layers surrounded by the connective tissue are reported: circular muscles forming a broad continuous band, longitudinal muscle bundles adjacent to the circular muscles, and dilator muscles crossing the connective tissue vertically toward the epithelium. The connective tissue has rosette glands. The esophagus of the larvae have epithelial cells with big vesicles but poorly developed interdigitations and filamentous structures, the cuticle is formed by a procuticle without differentiated exocuticle and endocuticle, the connective layer is thin and the rosette glands are absent. The observed features can be explained by his role in the swallowing of the food.     

A putative new hydrostatic skeletal function for the epidermis in Monhysterids (Nematoda)

The ultrastructure of the epidermis of two Monhysterid nematodes (Geomonhystera disjuncta and Diplolaimella dievengatensis) is studied in detail. The epidermis is composed of discrete uninucleated cells. The cytoplasmic layer of the epidermis between the cuticle and the somatic muscles is very thin and contains bundles of filaments that attach the muscles to the cuticle. The epidermal chords are voluminous and contain the nuclei and most of the cell organelles. In the chords many large electron-transparent vacuoles are found. It is hypothesized that these vacuoles fulfill a function as a compartmentalised hydrostatic skeleton.     

The fine structure of muscle attachments in an acarid mite, Caloglyphus mycophagus (Megnin) (Acarina)

The fine structure of the myo-cuticular junction in an acarid mite, Caloglyphus mycophagus, is described. The muscle fibres are attached to the cuticle via flattened, much invaginated, epidermal cells. Unlike the situation described for other arthropods, the stress across these epidermal cells does not appear to be transmitted by microtubules but rather by desmosome-like structures which form intraepidermal cell bridges where invaginations from the outer and inner surfaces of the epidermal cells lie close together. The muscles are attached to the inner surface of this complex desmosome and the outer surface is linked to the cuticle by extracellular fibrils.     

Anatomy of external structure and musculature of abdominal segments in the larva of the lepidopteran Pieris rapae crucivora

The external structure of the 1st (AS1) and 4th abdominal segments (AS4) of Pieris rapae is described in terms of pattern of shallow grooves on the cuticle. Both segments have 5 dorsal costae, 3 ventral costae, and an antero-posterior line in addiction to the dorsal and ventral intersegmental folds and a spiracle. AS4 has a pair of prolegs. The musculatures of AS1 and AS4 consist of 44 and 51 muscles, respectively. As in thoracic ones, most attachments of the muscles are located on the cuticular grooves. AS1 and AS4 have similar musculatures. Common to both segments are 89% of AS1 muscles and 84% of AS4 muscles. AS1 has 6 muscles homologous to proleg ones of AS4, including proleg retractors and plantar retractors. Comparison of the musculature of proleg-bearing abdominal segments among different species shows that abdominal musculature of lepidopteran larvae has major homologous and minor specific muscles. From the muscle attachment sites, the role of each muscle is inferred for contraction and bending of the body, lifting up its venter, taking off the crockets from the substrate, and retraction, lateral abduction, and anterior movement of the proleg.     

First Record of Soft Tissue Preservation in the Upper Devonian of Poland

Soft tissue preservation is reported from Upper Devonian deposits of the Holy Cross Mountains, central Poland, for the first time. The preserved soft tissues are muscles associated with arthropod cuticle fragments. The muscles are phosphatized with variable states of preservation. Well-preserved specimens display the typical banding of striated muscles. Other muscle fragments are highly degraded and/or recrystallized such that their microstructure is barely visible. The phosphatized muscles and associated cuticle are fragmented, occur in patches and some are scattered on the bedding plane. Due to the state of preservation and the lack of diagnostic features, the cuticle identification is problematic; however, it may have belonged to a phyllocarid crustacean. Taphonomic features of the remains indicate that they do not represent fossilized fecal matter (coprolite) but may represent a regurgitate, but the hypothesis is difficult to test. Most probably they represent the leftover remains after arthropod or fish scavenging. The present study shows that soft tissues, which even earlier were manipulated by scavenger, may be preserved if only special microenvironmental conditions within and around the animal remains are established.     

Exoskeleton anchoring to tendon cells and muscles in molting isopod crustaceans

Specialized mechanical connection between exoskeleton and underlying muscles in arthropods is a complex network of interconnected matrix constituents, junctions and associated cytoskeletal elements, which provides prominent mechanical attachment of the epidermis to the cuticle and transmits muscle tensions to the exoskeleton. This linkage involves anchoring of the complex extracellular matrix composing the cuticle to the apical membrane of tendon cells and linking of tendon cells to muscles basally. The ultrastructural arhitecture of these attachment complexes during molting is an important issue in relation to integument integrity maintenance in the course of cuticle replacement and in relation to movement ability. The aim of this work was to determine the ultrastructural organization of exoskeleton - muscles attachment complexes in the molting terrestrial isopod crustaceans, in the stage when integumental epithelium is covered by both, the newly forming cuticle and the old detached cuticle. We show that the old exoskeleton is extensively mechanically connected to the underlying epithelium in the regions of muscle attachment sites by massive arrays of fibers in adult premolt Ligia italica and in prehatching embryos and premolt marsupial mancas of Porcellio scaber. Fibers expand from the tendon cells, traverse the new cuticle and ecdysal space and protrude into the distal layers of the detached cuticle. They likely serve as final anchoring sites before exuviation and may be involved in animal movements in this stage. Tendon cells in the prehatching embryo and in marsupial mancas display a substantial apicobasally oriented transcellular arrays of microtubules, evidently engaged in myotendinous junctions and in apical anchoring of the cuticular matrix. The structural framework of musculoskeletal linkage is basically established in described intramarsupial developmental stages, suggesting its involvement in animal motility within the marsupium.     

Morphology and physiology of a hair plate sensory organ located on the antenna of the rock lobster Palinurus vulgaris

At the level of the J1 joint of each antenna of the rock lobster Palinurus vulgaris a hair plate sensory organ (hp) similar to those described in insects has been observed. The hp is located on the internal side of the S2 segment of the antenna, close to the soft articulating membrane of the J1 joint. It is formed by a triangular surface of cuticle about 3mm2 in area, covered with numerous hairs of different lengths (Figs. 1 and 2). Details of the hp were studied by scanning electron microscopy (Fig. 2). Physiological stimulation of the hp hairs occurs during medial movement of the J1 joint. Under this condition the soft articulating membrane rolls over the hairs and bends them progressively back onto the cuticle. Flexion of all the hairs corresponds to a medial movement of the J1 through 40 degrees. During this type of movement, the number of successively flexed hairs increases linearly (Fig. 3). Electrophysiological recordings of the hp sensory nerve correlated with selective mechanical stimulation of individual hairs demonstrated that each hair is innervated by a single sensory fiber. This sensory neurone responds phasically when the hair is flexed back onto the cuticle (as during an S2 medial movement) and when it returns to its resting position (as during an S2 lateral movement). Most of the sensory neurones are sensitive to the movement velocity of the hairs (Figs. 4 and 5). When the hair is maintained flexed its sensory neurone discharges tonically (Fig. 4). Electrical stimulation of the hp sensory nerve induced reflex actions in the external and internal rotator muscles of the segment S1. These effects were found to selectively activate the tonic motor command of these muscles (Fig. 6).     

The female abdomen of the viviparous earwig Hemimerus vosseleri (Insecta: Dermaptera: Hemimeridae), with a discussion of the postgenital abdomen of Insecta

The viviparous, epizoic African earwigs of the genus Hemimerus are currently regarded as the sister taxon of the remaining Dermaptera (Forficulina). Exoskeleton, musculature, and part of the nervous system of the female abdomen, from segment IV on, are described. The morphological interpretation and homology relations of most components are discussed, using previous and original data on Forficulina, Zygentoma, Ephemeroptera, Orthoptera and Dictyoptera as a comparative framework. In the mid-abdominal segments some interesting similarities with Zygentoma are indicated. Focal issues in the postgenital abdomen are the terminal dorsal sclerites, the cercal muscles, and the paraprocts and associated muscles. Earlier hypotheses on the dermapteran postabdomen (opisthomere and pseudocercus hypotheses) and results from ontogenetic studies are scrutinized. Some interesting features detected in female Hemimerus are the immobilization of terga VIII-X by means of a thick internal cuticle layer, the lack of dorsal muscles on these terga, the shift of some insertions of cercal and rectal muscles from tergum X to tergum IX, and minute pits on the venters IX and X that could be spiracle vestiges. Some of these features occur also in other Dermaptera. Some abdominal characters suggest that Hemimerus is nested within the Forficulina. The lack of the clasper-shape in the cerci is not a strong argument against this.     

A fine structural analysis of the epidermis and cuticle of the oligochaete aeolosoma Bengalense stephenson

The fine structure of the epidermis and cuticle has been described for the oligochaete Aeolosoma bengalense. The epidermis is a pseudostratified epithelium and consists of the following cell types: ciliated and nonciliated supportive cells, pigment cells and associated satellite cells, mucous cells, basal cells, and ciliated non-supportive columnar cells. Overlying and restricted to the supportive cells is a delicate cuticle composed of: (a) a discontinuous layer of membrane-bounded surface particles; (b) a thin filamentous layer of moderate electron density just under the surface particles; (c) a thicker inner filamentous layer of low electron density. Digestion with pronase effectively removes the cuticle. This, together with the fact that it stains with alcian blue and ruthenium red, indicates that the cuticle contains an acid mucopolysaccharide. Regeneration of the cuticle, following pronase treatment, is marked by the elaboration of numerous microvilli by the supportive cells. Most of the microvilli are transitory and evidence supports a microvillar origin for the cuticular surface particles. The presence of cuticular surface particles may be a characteristic shared in common by all oligochaetes and, perhaps, some polychaetes.     

On the pronymphal stage in the migratory locust (Locusta migratoria, Orthoptera, Acrididae)

The structure of the integument, somatic and visceral muscles, midgut, and Malpighian tubules were investigated at the late stages of the embryonic and early postembryonic development of the migratory locust, Locusta migratoria, to assess the organization of its pronymphal stage. In its morphogenetic features, the vermiform locust larva sometimes called the pronymph corresponds to the first nymphal instar covered with the second embryonic cuticle which has not been shed. Since the first-instar locust nymphs before and after the shedding of this embryonic cuticle differ significantly in many morphological characters, two consecutive phases of this nymphal instar can be distinguished: the first phase existing from the moment of development of the third embryonic cuticle to the shedding of the second one; the second phase existing from the shedding of the second embryonic cuticle to the formation of the cuticle of the second nymphal instar. Since the pronymphal stage should precede the nymph stage, it may be concluded that the pronymph of the locust is fully embryonized and covered with the second embryonic cuticle, which is also typical of other insects with hemimetabolous development (Konopová and Zrzavý, 2005). Therefore, it would be erroneous to refer to the vermiform first-instar nymph as the pronymph, because the two stages are separated by molting and formation of a new cuticle.     

Functional organization of the arctiid moth tymbal (insecta,lepidoptera)

The exoskeletal morphology, muscular organization, and innervation patterns of the tymbals of seven sound-producing species of tiger moths (Arctiidae) were compared with the undifferentiated episterna of two silent species. At least three muscles are involved in sound production: the tymbal muscle, pv2, and the accessory muscles, pvl and/or pv6. All of the tymbal muscles are innervated by the IIIN2a branch of the metathoracic leg nerve, which contains two axons larger than the others. Backfills of the tymbal branch of the IIIN2a reveal a medial sensory neuropil and a population of five ipsilateral motor neurons whose somata are clustered into three groups along the anterior edge of the metathoracic ganglion. The dendritic arborizations of the motor neurons extend to the ganglionic midline but are separate from one part of the auditory neuropil observed in other noctuoids. The study concludes that the arctiid tymbal reveals only minor modifications (e.g., cuticle thinning) of the episterna of silent moths and represents a primitive form of the tymbal compared to those of the Cicadidae.     

Electron microscopical localization of chitin in the cuticle of Halicryptus spinulosus and Priapulus caudatus (Priapulida) using gold-labelled wheat germ agglutinin: phylogenetic implications for the evolution of the cuticle within the Nemathelminthes

 The ultrastructure of the cuticle of adult and larval Priapulus caudatus and Halicryptus spinulosus is investigated and new features of cuticle formation during moulting are described. For the localization of chitin by TEM wheat germ agglutinin coupled to colloidal gold was used as a marker. Proteinaceous layers of the cuticle are revealed by digestion with pronase. The cuticle of larval and adult specimens of both species consists of three main layers: the outer, very thin, electron-dense epicuticle, the electron-dense exocuticle and the fibrillar, electron-lucent endocuticle. Depending on the body region, the exocuticle comprises two or three sublayers. The endocuticle can be subdivided into two sublayers as well. In strengthened parts such as the teeth, the endocuticle becomes sclerotized and appears electron-dense. Only all endocuticular layers show an intense labelling with wheat germ agglutinin-gold conjugates in all investigated specimens. Additional weak labelling is observed in the exocuticle III layer of the larval lorica of P. caudatus. All other cuticular layers remain unlabelled. Chitinase dissolves the unsclerotized endocuticular layers almost completely, but also exocuticle II and partly the loricate exocuticle III. The epicuticle, the homogeneous exocuticle I and the sclerotized endocuticle are not affected by chitinase. The labelling is completely prevented in all layers after incubation with chitinase. Pronase dissolves all exocuticular layers, but not evenly. The presumably sclerotized regions of exocuticle I are not affected as well as the complete epicuticle and the endocuticle. All cuticular features of the Priapulida are compared with the cuticle of each high-ranked taxon within the Nemathelminthes with special regard to the occurrence of chitin. Based on this out-group comparison it can be concluded that: (1) a two-layered cuticle with a trilaminate epicuticle and a proteinaceous basal layer represents an autapomorphic feature of the Nemathelminthes, (2) the stem species of the Cycloneuralia have already evolved an additional basal chitinous layer, (3) such a three-layered cuticle is maintained as a plesiomophy in the ground pattern of the Scalidophora and (4) in the Nematoida, the chitinous basal layer is replaced by a collagenous one at least in the adults; the synthesis of chitin is restricted to early developmental phases or the pharyngeal cuticle. Accepted: 12 March 1998     

Fine Structure of the Archiannelid Cuticle and Remarks on the Evolution of the Cuticle Within the Spiralia*

The fine structure of the cuticle in the archiannelids Polygordius, Protodrilus, Trilobodrilus, and Diurodrilus is described. Polygordius and Protodrilus are unique with respect to the filiform microvilli penetrating through the cuticle. Basically all forms conform to the most common type of cuticular structure in the Annelida: a fibrous matrix of varying complexity penetrated by micro-villar-like cell processes. Following the ideas expressed by Lyons (1970), a hypothesis on the evolution of the cuticle within the vermiform Spiralia (e.g. Turbellaria, Nemertea, Annelida) is proposed. New data on interstitial Turbellaria and other new forms of interstitial worms suggest a morphological and functional sequence from the multiciliated epidermis of Turbellaria to the cuticularized hypodermis of Annelida.     

Metamorphic changes of wild type and mutant Drosophila tissues induced by 20-hydroxy ecdysone in vitro

Wild type (Oregon R) and non-pupariating as well as late-pupariating mutant larval tissues were cultured in vitro up to 5 weeks with and without 20-hydroxy ecdysone (1 μg/ml). The following responses were elicited by the hormone: in the case of wild type tissues detachment of the larval epidermis and muscles from the cuticle; puparial tanning and sclerotization of the larval cuticle; dissociation of the fat body into single cells; inhibition of the movement of the hind intestine. Most of these responses developed within 1 week of culturing. Of the 4 mutants tested, 3 behaved like the wild type. In cultures of ?(1)npr-1, however, puparial tanning, disc evagination, and inhibition of the movement of the hind intestine was abnormally weak and the dissociation of fat body was not observed at all. Detachment of the epidermis and muscles as well as formation of the pupal cuticle by disc tissue occurred normally. The results are discussed with respect to the ecdysteroid-induced metamorphosis of the tissues and the autonomy of mutant gene action.     

Changes in the first instar cuticle of Schistocerca gregaria before and associated with hatching

Size increases in the first instar cuticle are described in the pharate stage, at hatching and ecdysis, and in the hour following ecdysis. The ways in which the cuticle expands are described. Changes in colour and the mechanical properties of cuticle are described and related to changes in the different protein fractions extracted from the cuticle. A blood-borne factor is shown to affect darkening in the first instar.     

Structure of the stomach cuticle in adult and larvae of the spider crab Maja brachydactyla (Brachyura,Decapoda)

The stomach of decapods is a complex organ with specialized structures that are delimited by a cuticle. The morphology and ontogeny of the stomach are largely described, but few studies have focused on the morphology of its cuticle. This study examined the morphology of the stomach cuticle of cardiac sacs, gastric mill ossicles, cardio-pyloric valve and pyloric filters, and during various stages (zoea I and II, megalopa, first juvenile, and adult) of the common spider crab Maja brachydactyla using dissection, histology and transmission electron microscopy. The results show that cuticle morphology varies among structures (e.g., cardiac sacs, urocardiac ossicle, cardio-pyloric valve, pyloric filters), within a single structure (e.g., different sides of the urocardiac ossicle) and among different life stages. The cuticle during the larval stages is very thin and the different layers (epicuticle, exocuticle, and endocuticle) are infrequently distinguishable by histology. Major changes during larval development regarding cuticle morphology are observed after the molt to megalopa, including the increment in thickness in the gastric mill ossicles and cardio-pyloric valve, and the disappearance of the long thickened setae of the cardio-pyloric valve. The cuticle of all the stomach structures in the adults is thicker than in larval and juvenile stages. The cuticle varies in thickness, differential staining affinity and morphology of the cuticle layers. The structure–function relationship of the cuticle morphology is discussed.     

The structure of developing muscle insertions in insects

The relationship between muscles and the components of the integument in muscle insertions have been studied with the electron microscope in two insects, Calpodes ethlius (Hesperiidae, Lepidoptera) and Rhodnius prolixus (Reduviidae, Hemiptera). The area of contact between the muscles and the epidermis is increased by interdigitating processes whose membranes are joined by intermediate junctions. The junctions occur at the level of a Z line so that actin filaments attach directly to them. Within the epidermis, microtubules extend from the junctions of the myoepidermal connection to the cuticle, where they attach to hemidesmosomes which line deep indentations of the membrane. The microtubules probably enable the tendinous epidermal cells to withstand the tensions exerted upon them by the muscles. The epidermis is anchored to the cuticle by tonofibrillae, homogeneous rods secreted in the deep indentations of the plasma membrane. Since the tonofibrillae of successive instars are continous, they penetrate and attach to the cuticulin, the outermost layer of the epicuticle.     

Structure, function and evolution of somatic musculature in Dasydytidae (Paucitubulatina, Gastrotricha)

The musculature of two species of the gastrotrich taxon Dasydytidae, Dasydytes (Dasydytes) goniathrix and Haltidytes crassus, was investigated and described using phalloidin staining, confocal microscopy and computer-aided three-dimensional data analysis. Dasydytidae is a peculiar taxon of freshwater Gastrotricha, containing species that are characterized by different adaptations to a semiplanktonic lifestyle, a rather uncommon feature among primarily benthic Gastrotricha. Like other dasydytid species studied so far, D. goniathrix and H. crassus possess a system of movable cuticular spines with an associated system of somatic oblique and segmented lateral muscles. The presence of other somatic (dorsodermal muscles R1 and R2) and visceral muscles (musculi ventrales, m. ventrolaterales, m. dorsales, m. helicoidales) known from a wide range of gastrotrich species was confirmed. Regarded from a functional perspective, the earlier proposed antagonistic role of oblique muscles (as spine abductors) and segmented lateral muscles (as adductors) is questioned for the species studied herein. Alternatively, our structural and behavioral observations suggest that muscular spine abduction in D. goniathrix is brought about by synergistic contraction of the musculi obliqua and m. laterales, and a passive adduction due to muscle relaxation and elastic recoil of the trunk and cuticle. For H. crassus, we hypothesize active muscular spine abduction by contraction of the musculi obliqua plus the last segment of m. laterales accompanied by severe cuticle deformations close to the spine insertions. Adduction is achieved by cuticle reformation due to elasticity and increase in tissue pressure brought about by muscle action, possibly of enforced dorsodermal muscles. The newly obtained and published muscular data of further gastrotrich species were gathered in a species-character matrix. Based on this data set, a maximum parsimony analysis of representatives of the Dasydytidae has been conducted. According to this analysis, there are three well-supported monophyletic lineages within likewise monophyletic Dasydytidae. The first lineage comprises the taxa Anacanthoderma, Stylochaeta and Chitonodytes, the second comprises Dasydytes, Setopus and Ornamentula, and the third represents the taxon Haltidytes. Relationships between these clades could be resolved but are only weakly supported. The new phylogenetic hypothesis is used to reconstruct the ancestral character pattern and to infer possible evolutionary transformations within the Dasydytidae.