Ultrastructure of multicellular foregut glands in selected Solenogastres (Mollusca)

Foregut glands in Solenogastres are not only of importance in biological processes like feeding and digestion, but multicellular foregut glands also provide valuable characters for taxonomy and systematics. Here, the fine structure of four different types of foregut glands is investigated: the Meioherpia-type ventrolateral foregut gland of Meioherpia atlantica, the Simrothiella-type ventrolateral foregut gland of Simrothiella cf. margaritacea, and the Pararrhopalia-type ventrolateral foregut gland as well as the dorsal gland of Pararrhopalia pruvoti. Thereby, special focus is set on the arrangement of glandular and supporting cells within the glands, and on the characterization of glandular cells according to the size, shape and electron-density of their secretional vesicles. It is shown that the investigated glands are complex organs composed of non-glandular supporting cells and two to five glandular cell types producing discrete secretions.     

Ultrastructure and histochemistry of the foregut in<Emphasis Type="Italic"> Wirenia argentea</Emphasis> and<Emphasis Type="Italic"> Genitoconia rosea</Emphasis> (Mollusca,Solenogastres)

Wirenia argentea and Genitoconia rosea feed on Cnidaria like most representatives of the molluscan taxon Solenogastres (Aplacophora, Neomeniomorpha sensu Scheltema). The structure and histochemistry of the foregut are described based on histologic, semithin, and ultrathin section series. The ultrastructure was analyzed by means of transmission electron microscopy. There are two sets of unicellular glands: a narrow row of preoral gland cells opening to the preoral area, and pharyngeal gland cells in high numbers. Preoral gland cells produce serous secretions in W. argentea, but mucosubstances in G. rosea, whereas pharyngeal gland cells are similar in structure and histochemistry in both species. Based on the size and electron density of gland vesicles, five distinct types of pharyngeal gland cells can be defined. In contrast to earlier assumptions, all types of pharyngeal gland cells produce serous secretions, most probably representing digestive ferments, but no mucosubstances.     

The digestive tract of Helicoradomenia (Solenogastres, Mollusca), aplacophoran molluscs from the hydrothermal vents of the East Pacific Rise

Abstract. Species of Helicoradomenia are constantly found at hydrothermal vent sites of the eastern and western Pacific Ocean. The digestive tract of 2 species of the genus was investigated with special focus on the ultrastructure and histochemistry of epithelia and glandular organs. The preoral cavity and foregut epithelia are composed of microvillous main cells, secretory cells producing protein-rich substances, and sensory cells with specialized cilia. The foregut bears a pair of glands with 3 types of extremely long-necked glandular cells surrounded by musculature. Each glandular cell opens directly into the radula pocket without a gland duct. The large radula apparatus consists of pairs of denticulated bars resting on a flexible radular membrane without elaboration of a subradular membrane. The midgut has a narrow, mid-dorsal tract of ciliary cells, but most of the epithelium is composed of digestive cells with a highly developed lysosomal system. The hindgut is lined by ciliated cells and free of glands. The foregut and radula seem to be highly efficient in the capture of relatively large, motile prey. Food contents within the midgut lumen and within some of the large secondary lysosomes indicate a triploblastic metazoan prey of non-cnidarian origin. The digestive tract is not adapted to microvory and there is no indication of a symbiosis with chemoautotrophic bacteria.     

Ultrastructural investigation of a salivary gland in a centipede: structure and origin of the maxilla I-gland of Scutigera coleoptrata (Chilopoda, Notostigmophora)

The maxilla I-gland of Scutigera coleoptrata was investigated using light and electron microscopy methods. This is the first ultrastructural investigation of a salivary gland in Chilopoda. The paired gland opens via the hypopharynx into the foregut and extends up to the third trunk segment. The gland is of irregular shape and consists of numerous acini consisting of several gland units. The secretion is released into an arborescent duct system. Each acinus consists of multiple of glandular units. The units are composed of three cell types: secretory cells, a single intermediary cell, and canal cells. The pear-shaped secretory cell is invaginated distally, forming an extracellular reservoir lined with microvilli, into which the secretion is released. The intermediary cell forms a conducting canal and connects the secretory cell with the canal cell. Proximally, the intermediary cell bears microvilli, whereas the distal part is covered with a distinct cuticle. The cuticle is a continuation of the cuticle of the canal cells. This investigation shows that the structure of the glandular units of the salivary maxilla I-gland is comparable to that of the glandular units of epidermal glands. Thus, it is likely that in Chilopoda salivary glands and epidermal glands share the same ground pattern. It is likely that in compound acinar glands a multiplication of secretory and duct cells has taken place, whereas the number of intermediary cells remains constant. The increase in the number of salivary acini leads to a shifting of the secretory elements away from the epidermis, deep into the head. Comparative investigations of the different head glands provide important characters for the reconstruction of myriapod phylogeny and the relationships of Myriapoda and Hexapoda.     

Exocrine glands in the legs of the social wasp Vespula vulgaris

This study brings a survey of the exocrine glands in the legs of Vespula vulgaris wasps. We studied workers, males, virgin queens as well as mated queens. A variety of 17 glands is found in the different leg segments. Among these, five glands are novel exocrine structures for social insects (trochanter-femur gland, ventrodistal tibial gland, distal tibial sac gland, ventral tibial gland, and ventral tarsomere gland). Most leg glands are present in the three leg pairs of all castes. This may indicate a mechanical function. This is likely for the numerous glands that occur near the articulation between the various leg segments, where lubricant production may be expected. Other possible functions include antenna cleaning, acting as a hydraulic system, or pheromonal. Further research including leg-related behavioural observations and chemical analyses may help to clarify the functions of these glandular structures in the legs.     

Molluscan muscle systems in development and evolution*

The evolutionary history of the various molluscan muscle systems reflects drastic modifications and reductions as well as true innovations. No less than eight main and independent muscle systems of the Mollusca are described and, based on the current understanding of molluscan phylogeny, their evolutionary histories are outlined.New data on the myogenesis of the Polyplacophora by means of fluorescence‐staining and image analysis by confocal laser scanning microscopy show that the pre‐oral region recapitulates a ‘worm‐grid’, and that the dorso‐ventral musculature passes a stage of multiple seriality as found in adult Solenogastres. Old and new data on bivalves and recent studies on primitive gastropods provide clear evidence that the larval musculature of both groups (and thus possibly of all conchiferans) is entirely independent from the adult condition. The growth of shell‐inserted muscles always necessitates substantial renewal of myocytes which is still poorly understood. Although very promising for phylogenetic purposes, the understanding of the developmental genetics of the various molluscan muscle systems is still in its infancy.     

Gill‐derived glands in species of Astyanax (Teleostei: Characidae)

The presence of a gill‐derived gland is herein reported for the first time in males of species of Astyanax and related genera; they are described through histological cuts and SEM. The gill‐derived glands described for the Characidae, when fully developed, present a similar structure in different species. The main external feature of gill‐derived glands is the fusion of anteriormost gill filaments on the ventral branch of first gill arch. This fusion is caused by squamous stratified epithelial tissue that covers adjacent filaments, forming a series of chambers. In the region where the gill‐derived gland develops, the secondary lamellae of the gill filaments are much reduced or completely atrophied being characterized by the presence of glandular cells forming nests.     

Morphology and ultrastructure of the pharynx in Solenofilomorphidae (Acoela)

The homology of pharynges within the mostly pharynx-less Acoela has been a matter of discussion for decades. Here, we analyze the pharynges of three members of the Solenofilomorphidae, Myopea sp. and two species of the genus Solenofilomorpha, by means of light and transmission electron microscopy. Special focus is placed on the ultrastructure of the pharyngeal musculature, epidermis surrounding the mouth, pharyngeal epithelium, and junction with the digestive parenchyma. The main goal of this study was to evaluate the usefulness of certain characters for broader comparisons within the Acoela. Among the three species, characters relating to position of the mouth, presence and elaboration of sphincter muscles, presence of pharyngeal glands, and ultrastructure of epitheliosomes proved to be variously species- and genus-specific. The arrangement of pharyngeal muscles and their connection with body wall musculature, ultrastructure of receptor cells, and morphology of a nonciliated glandular region in the posterior pharynx, in contrast, appear to be characteristic of the family Solenofilomorphidae and thus of predominant interest for comparisons with other acoel families.     

Ultrastructure of the labrum and foregut of Derocheilocaris remanei (Crustacea,Mystacocarida)

The cuticle-lined foregut of Derocheilocaris remanei consists of the mouth with its associated labrum, and an undifferentiated esophagus. It is separated from the midgut by an esophageal valve. The labrum is a conspicuous structure moved by five pairs of muscles (four dorsoventral and one longitudinal). Four pairs of subcuticular glands open to its inner face forming two longitudinal, lateral rows of cuticular pores. Each secretory unit is composed of a glandular component (with one or two secretory cells), a neck cell, and a duct cell. In addition, a single gland cell opens mesially into the buccal cavity. The ventrally located mouth is a complex structure characterized by a filter-like system, a sensory organ, and epithelial cells with highly developed microvilli. The esophagus is a simple tube with a characteristic curvature following the mouth. It has a rounded cross section and a triradiate lumen. A layer of circular musculature surrounds this region. The end of the esophagus protrudes into the midgut lumen forming the so-called esophageal valve. The ultrastructural features of the foregut, with the presence of a mucus-trapping mechanism, a relatively well-developed filter system and associated structures and an esophagus lacking glands confirm the microphagic feeding habits of mystacocarids. © 1996 Wiley-Liss, Inc.     

Immunocytochemical localization of seminal proteins in salivary and lacrimal glands of the rat

Antibodies against 10 different secretory proteins from the accessory sex glands of the male rat were used for immunohistochemical studies of salivary and lacrimal glands from intact and castrated rats, at the light- and electron-microscopic levels. In the parotid gland, secretory acinar cells showed immunoreactivity with antibodies against prostatic binding protein, cystatin-related peptide and acid phosphatase (isoenzyme pI 8.0; 5.6) typical of ventral prostate, and seminal vesicle secretion VI. Western blotting analysis indicated that immunoreactivity against prostatic binding protein was attributable to a subunit, presumably C3. Acid phosphatase pI 5.6 showed a molecular weight of 66 kDa, which is at variance with the prostatic form. Immunoreactivity for secretory transglutaminase, derived from the coagulating gland, was restricted to myoepithelial and stromal cells. In castrated animals, the immunoreactivity of acinar cells was reduced to the background level, whereas stromal transglutaminase immunoreactivity was unaltered. The distribution pattern of immunoreactivity for the proteins mentioned was almost identical in the lacrimal gland. Significant differences were however observed in the immunoreactivity of the inframandibular gland, where serous glandular cells were non-immunoreactive for seminal proteins, with the exception of acid phosphatase isoenzyme pI 8.0. Granules present in the convoluted granular ducts were immunoreactive particularly for acid phosphatase (isoenzyme pI 5.6)but much less for cystatin-related peptide; immunoreactivity was reduced after castration. The straight portion of the inframandibular duct system was immunoreactive for transglutaminase, but no influence of castration was visible. The distribution of immunoreactivity for seminal proteins present in the salivary and lacrimal glands and the pronounced androgen-dependence of their expression point to functional relationships of the respective proteins at both glandular sites.     

The development of lingual glands in the domestic duck (Anas platyrhynchos f. domestica): 3D‐reconstruction,LM, and SEM study

The major salivary glands of birds develop by branching or elongation of the epithelial cords. The development of the minor salivary glands in form of the lingual glands has never been described. Among birds, only Anatidae have three types of the lingual glands: rostral, caudo‐lateral, and caudo‐medial lingual glands. The study aims to characterize the manner and rate of the lingual glands development in the domestic duck and their topographical arrangement relative to the hyoid apparatus. The study reveals that all three types of the lingual glands develop by branching. We describe five stages of the lingual glands development in the domestic ducks: prebud, initial bud, pseudoglandular, canalicular, and terminal bud stage. The pattern of the lingual glands development in birds is similar to that described for mammals, with the exception, that the terminal buds are formed at the same time as the lumen of the glands. Generally, the rostral lingual gland starts to branch earlier than the caudal lingual glands. The 3D‐reconstruction shows the location and direction of lingual gland development relative to the entoglossal cartilage and basibranchial bone. Light microscopy and scanning electron microscopy allow to characterize the histogenesis of the embryonic epithelium into glandular epithelium. At a time of hatching only secretory units of caudal lingual glands resemble the secretory units of the adult domestic duck. The rostral and caudo‐lateral lingual glands are arranged on the sides of the entoglossal cartilage and basibranchial bone and caudo‐madial lingual glands are located over the basibranchial bone. We suggest that such an arrangement of the lingual glands in the domestic duck is important during food intake and responsible for reduction of friction and formation of food bites.     

Morphology of the salivary glands of three Squamata species: Podarcis sicula sicula, Tarentola mauritanica and Coluber viridiflavus

The histology, histochemistry and ultrastructure of the salivary glands of three species of squamata, Podarcis sicula sicula (mandibular glands), Tarentola mauritanica (sublingual gland) and Coluber viridiflavus (supra- and infralabial glands) were studied. Each gland contained acidic cells, positive for both periodic acid Schiff and Alcian blue reactions. These cells can be distinguished as seromucous and mucous types based on the different electron density of their granules. α-Amylase, until now detected only in mammalian salivary glands, was not found in any of the salivary glands examined. The ultrastructural study revealed that the salivary glandular cells of T. mauritanica lack intercellular canaliculi, which by contrast, are present between the seromucous cells in the salivary glands of C. viridiflavus and P. s. sicula . Comparable variation is also seen when the ultrastructural features of the secretory granules in salivary glands of the three Squamata species are compared. The salivary granules of T. mauritanica and C. viridiflavus are more or less dense but structureless, while the mucous granules of P. s. sicula have a distinctive and characteristic substructure. Therefore, this study, designed to obtain comparative data on the histology, histochemistry and ultrastructure of the salivary glands of three representative, but hitherto unstudied, species of Squamata, reveals great variation in the structure of these glands within the Squamata lineage, even when compared to previously documented species.     

Opisthonotal glands in the Camisiidae (Acari, Oribatida): evidence for a regressive evolutionary trend

Paired, sac-like and typically large opisthonotal glands (syn. oil glands), mainly considered for chemical protection and communication, characterize the so-called 'glandulate Oribatida' which include the Parhyposomata, Mixonomata, Desmonomata and Brachypylina but also the Astigmata. Among these groups distinct evolutionary trends affect the morphology of glands and their secretion profiles, thereby rendering them highly informative characters with phylogenetic significance. One striking tendency, convergently occurring in a few glandulate groups, leads to the degeneration or even complete regression of opisthonotal glands. In this study, a first example of coherent evolutionary steps towards opisthonotal gland degeneration is described by using desmonomatan Camisiidae as a model: Opisthonotal glands in representatives of genus Platynothrus still show morphologically and chemically ancient conditions with fairly-well developed glandular reservoirs. Secretion patterns mainly consist of a characteristic set of terpenes and aromatics ('astigmatid compounds') as found in outgroups such as desmonomatan Trhypochthoniidae. Progressive states of regression of opisthonotal glands, along with a reduction of component-richness and amounts of secretions, occur in representatives of Heminothrus and, more conspicuously, in species of Camisia , most likely indicating a consistent evolutionary trend. This trend towards opisthonotal gland atrophy may be due to novel alternative and cheap strategies of passive defense in more-derivative camisiids – such as mechanical protection by encrustation of the cuticle – that possibly compensate for the lack of chemical defenses.     

Parapodial glandular organs in Spiophanes (Polychaeta: Spionidae)—studies on their functional anatomy and ultrastructure

Parapodial glandular organs (PGOs) of Spiophanes (Polychaeta: Spionidae) were studied using light and electron microscopy. These organs are found in parapodia of the mid body region, starting on chaetiger 5 and terminating with the appearance of neuropodial hooks (chaetiger 14 or 15 in adult individuals). Large PGOs in anterior chaetigers display different species‐specific types of openings whereas small PGOs in posterior parapodia of the mid body region always open in a simple vertical slit. Each PGO is composed of three main complexes: (1) the glandular sac with several distinct epithelia of secretory cells and secretory cell complexes and the reservoir filled with fibrous material, (2) the gland‐associated chaetal complex (including the region of chaetoblasts and follicle cells, follicular canals, two chaetal collector canals, the combined conducting canal, the chaetal spreader including the opening of the glandular organ with associated type‐1 secretory cells, and the gland‐associated chaetae), and (3) a bilayered musculature surrounding the gland. A considerable number of different cell types are involved in the secretory activity, in the guidance of the gland‐associated chaetae, and in the final expulsion of the fibrous secretion at the opening slit. Among these different cell types the type‐5 secretory cells of the proximal glandular complex with their cup‐shaped microvilli emanating thick microfibrils into the lumen of the glandular sac are most conspicuous. Secretory cells with cup‐shaped microvilli being involved in the production of β‐chitin microfibrils have so far only been reported from some representatives of the deep‐sea inhabiting Siboglinidae (Polychaeta). We suggest that the gland‐associated chaetae emerging from inside the PGOs of Spiophanes are typical annelid chaetae formed by chaetoblasts and follicle cells. Functional morphology implies the crucial role of PGOs in tube construction. Furthermore, the PGOs are discussed in consideration of phylogenetic aspects. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Morphological analysis of the male reproductive accessory glands of the bat Artibeus lituratus (Phyllostomidae: Chiroptera)

Bats are distributed worldwide from tropical to temperate regions. Despite their wide geographical radiation and advances in studies using evolutionary approaches, aspects related to the reproduction of these animals remain poorly explored, especially those related to the male reproductive accessory glands (RAGs). Thus, the aim of this study was to analyze the morphophysiology of the male RAGs in the bat Artibeus lituratus. The RAGs in A. lituratus are composed of a compact intra‐abdominal glandular complex, consisting of the prostate with two prostatic regions (ventral and dorsal), plus Littre glands and a pair of extra‐abdominal bulbourethral glands. The ventral region of the prostate has an epithelium with variable morphology, due to its holocrine type of secretion. In contrast, the dorsal region has a typical cubic‐to‐columnar pseudostratified epithelium. Both regions contain two cell types, basal and secretory cells. Similar to the epithelial morphology, the secretion also varies, with the ventral region containing numerous PAS‐positive globular vesicles, whereas the dorsal region has a more fluid, hyaline and PAS‐negative secretion. Littre glands are dispersed in the connective tissue of the urethra, while the bulbourethral glands are located in the penile root, both glands with cubic‐to‐columnar pseudostratified epithelium and globular PAS‐positive secretion. The results demonstrate that the RAGs of A. lituratus are composed of two prostatic regions, ventral and dorsal, and urethral and bulbourethral glands, with no seminal vesicles. Each prostatic region has unique and distinctive characteristics, with the ventral region presenting an exclusive holocrine nature and the dorsal region having similarities to the ventral prostate of rodents.     

Histochemical and ultrastructural study on the innervation of the byssus glands of Mytilus galloprovincialis

The aluminium-formaldehyde (ALFA) histofluorescence method reveals an extensive plexus of brilliant greenish monoaminergic elements in the glandular zones of the Mytilus foot, while only scanty nerve fibres are acetylcholinesterase-positive. By electron microscopy, bundles of nerve fibres can be seen i) in close connection with the intrinsic musculature located in the connective septa among the glands, and ii) near the cell bodies and necks of all the byssus glands. The nerve fibres show varicosities containing three types of vesicles: small clear (50-60 nm), small granular (80-90 nm), and large granular (160-200 nm). The regions of close apposition between nerve terminals and muscle or gland cells generally do not show typical pre- or postsynaptic specializations. Along the pedal groove, mainly in the proximal two thirds of the foot, peripheral bipolar neurons can be detected, both by fluorescence and electron microscopy.     

Whale tear glands in the bowhead and the beluga whales: Source and function

Orbital glands are found in many tetrapod vertebrates, and are usually separate structures, consisting of individual glands lying in the eyelids and both canthi of the orbit. In cetaceans, however, the orbital glandular units are less distinct and have been described by numerous authors as a single, periorbital mass. There are few histochemical and immunhistochemical studies to date of these structures. In this study, we examined the orbital glandular region of both the bowhead whale (Balaena mysticetus: Mysticeti) and the beluga whale (Delphinapterus leucas: Odontoceti) using histological, histochemical, and immunohistochemical techniques. Histologically, in the bowhead, three glandular areas were noted (circumorbital, including Harderian and lacrimal poles), palpebral (midway in the lower eyelid), and rim (near the edge of the eyelid). In the beluga, there was only a large, continuous mass within the eyelid itself. Histochemical investigation suggests neither sexual dimorphism nor age-related differences, but both whales had two cell types freely intermingling with each other in all glandular masses. Large cells (cell type 1) were distended by four histochemically distinct intracellular secretory granules. Smaller cells (cell type 2) were not distended (fewer granules) and had two to three histochemically distinct intracellular secretory granules. The beluga orbital glands had additional lipid granules in cell type 1. Counterintuitively, both lipocalin and transferrin were localized to cell type 2 only. This intermingling of cell types is unusual for vertebrates in whom individual orbital glands usually have one cell type with one to two different secretory granules present. The heterogeneity of the orbital fluid produced by cetacean orbital glands implies a complex function, or series of functions, for these orbital glands and their role in producing the tear fluid.