Foregut glands of Solenogastres (mollusca): anatomy and revised terminology

In the molluscan class Solenogastres, different types of foregut glands vary in number, structure, and location within the foregut. The present article describes their anatomy and cytology and intends to clarify their confused terminology. Pharyngeal glands, esophageal glands, and the more complex dorsal and ventrolateral foregut glands can be distinguished. The ventrolateral foregut glands (ventral foregut glandular organs, ventral salivary glands of auct.), in the literature subdivided previously into four types, are revisited here in the context of current vertebrate gland terminology. The results of recent investigations are added to earlier ones, and a classification system for these multicellular glands is proposed. This system is based on cytological characters of glandular cells (intra- or extraepithelial), characters of the associated musculature (inner or outer musculature), location of the gland relative to the pharynx epithelium (endoepithelial or exoepithelial), characters of the gland openings (paired or unpaired), morphology of the gland duct (simple or branched), and some additional features like the arrangement of glandular cells along the gland ducts. Gross morphology and anatomy of ventrolateral foregut glands constitute useful taxonomic characters in determining higher taxa (family level), and finer details of the anatomy and cytology are useful in determining lower levels (genus and species). Possible pathways for the evolution of the different gland types of Solenogastres in relation to foregut glands present in the other molluscan clades are presented. The importance of ventrolateral foregut gland characters for phylogenetic considerations within the Solenogastres is discussed.     

Fine structure of reproductive glands in two primitive marine pulmonates (Basommatophora: Siphonariidae)

Within the clade Euthyneura the marine basommatophorans are particularly neglected. More morphological and molecular studies are needed because their phylogenetic relationships with other pulmonates remain unresolved. The present study examines the most conspicuous reproductive gland, the glandular complex in two marine limpets, Siphonaria capensis and S. serrata (Pulmonata: Basommatophora) at both gross and fine structural levels. These two sympatric species with different developmental modes were selected to compare the structure and function of this enormous glandular structure. In both S. capensis and S. serrata, the glandular complex shows an undifferentiated state composed of an acidophilic albumen gland and a basophilic mucous gland. The glands contain secretory cells and supporting cells (= ciliated cells) that are highly ciliated. When the histochemical properties of the glandular complex were compared with those of siphonariid egg masses (of each species) it could be established that the albumen gland was responsible for the production of perivitelline fluid whereas the mucous gland secreted substances that help in the assembly of mucous layers surrounding the egg capsules. We suggest that the presence of a single glandular complex comprised of two glands is the most primitive organization of reproductive glands in pulmonates. Furthermore, the histology, fine structure and histochemistry of these glands are very similar to those of the reproductive glands of opisthobranchs.     

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.     

The development of amphids and amphidial glands in adult Syngamus trachea (Nematoda: Syngamidae)

Nematode amphids are a pair of lateral cephalic sense organs, each comprising a group of sensory endings terminating in a cuticle-lined pit. In Syngamus trachea, a parasite of birds, each amphid is surrounded by two non-nervous supporting elements, a large gland cell basally and a smaller supporting cell anteriorly. The amphidial glands display high levels of secretory activity from five to six days postinfection. Secretory material is discharged through the lumen of the sense organ onto host tissue. The ultrastructure of amphids and amphidial glands has been investigated in newly moulted, immature and mature adults to trace the development of glandular activity and its effect on amphid-amphidial gland relationships. In newly moulted adults, the glands have very low levels of secretory activity and appear to act only as supporting cells to the amphids. As secretory activity increases, the gland cell membrane surrounding the sensory endings is elaborated into a reticulum which probably forms the secretory surface. In mature adults the amphid pit is swollen and filled with secretion; the sensory endings are relegated to the periphery of the lumen. It is suggested that amphidial glands develop from typical supporting cells, but acquire a new role possibly associated with parasite attachment.     

Generation gland morphology in cordylid lizards: An evolutionary perspective

To elucidate the functional significance of the three distinct types of generation glands that have been identified among cordylid lizards, we mapped gland type to the terminal taxa in the most recent phylogenetic tree for the Cordylidae. We used the phylogenetic programme Mesquite and applied the principle of parsimony to infer character states for the ancestral nodes in the tree. For those species where information on gland type was not available from the literature, we conducted a histological investigation of generation gland morphology, using standard histological techniques. We included two species of the sister family Gerrhosauridae in the analysis to serve as outgroups. In both Gerrhosaurus typicus and G. flavigularis, scales immediately anterior to the femoral pores displayed glandular activity, but differed from generation glands of cordylids in the absence of mature glandular generations. Among the cordylids investigated, we identified a fourth type of generation gland in Pseudocordylus subviridis, P. spinosus, and in the two Hemicordylus species, one where the glands consistently comprise of two mature glandular generations. In H. capensis, both single‐ and two‐layer type glands are present. Our reconstruction of ancestral character states suggests a minimum of six transformations from one gland type to another during the evolutionary history of the family. The reconstruction furthermore suggests that the single‐layer type gland reappeared at least once (in Hemicordylus) in the Cordylinae after having been lost. The reconstruction also unequivocally shows that the pit‐like multiple‐layer type gland evolved directly from the single‐layer type and not from the protruding multiple‐layer type. The two‐layer type gland appears to be an intermediary condition between the multiple‐layer and single‐layer types. The evolutionary transformation of generation gland type appears to be linked to changes in lifestyle and associated changes in degree of territoriality and the need for chemical communication. J. Morphol. 275:456–464, 2014. © 2013 Wiley Periodicals, Inc.     

Ultrastructural study of two glandular systems in the proboscidial glandular epithelium of Riseriellus occultus (Nemertea, Heteronemertea)

 Two different types of glandular system in the proboscidial epithelium of Riseriellus occultus have been investigated by transmission electron microscopy. As expected, most of the epithelial cells are glandular in nature. With regard to differences in the ultrastructure of these gland cells and in the formation and morphology of their secretory granules, we have categorized and described four types of gland cell, indicated as G₁, G₂, G₃, and G₄. Each gland cell has a completely intraepithelial body characterized by a prominent nucleus, developed rough endoplasmic reticulum, Golgi complexes, and numerous secretory granules at different stages of maturation. These four types of gland cell appear associated in pairs forming numerous glandular systems of two types (A, B). These glandular systems are restricted to the ventral surface of the proboscis and are scattered irregularly throughout its length. Each glandular system consists of two gland cells of different types. The gland cell necks in each glandular system extend together to the epithelial surface; they protrude onto this and form a papilla where they open in a common area. The epithelial supportive cells adjacent to the glandular systems have long, stout microvilli which have a core of tonofilaments. These tonofilaments gather into dense bundles which pass vertically through the supportive cells and attach to the extracellular matrix underlaying the cells by hemidesmosomes. Moreover, a single sensory process stands close to each papilla. The ultrastructural morphology of the type A glandular systems suggests that they have an adhesive function operating in a similar way to that of the duo-gland adhesive systems in other invertebrate groups, although they are not homologous with these. The spatial arrangement of the secreted products of the type B glandular systems suggests that these may contribute to increasing the grip of the proboscis on the prey. The secretory granules (=pseudocnids) of the type G₃ gland cells are very likely an autapomorphy of the Anopla, providing a character by which the relationships within the Nemertea can be evaluated. Accepted: 9 October 1997     

MORPHOGENESIS OF CAPITATE GLANDULAR HAIRS OF CANNABIS SATIVA (CANNABACEAE)

The glandular secretory system in Cannabis sativa L. (marihuana) consists of three types of capitate glandular hairs (termed bulbous, capitate-sessile, and capitate-stalked) distinguishable by their morphology, development, and physiology. These gland types occur together in greatest abundance and developmental complexity on the abaxial surface of bracts which ensheath the developing ovary. Bulbous and capitate-sessile glands are initiated on very young bract primordia and attain maturity during early stages of bract growth. Capitate-stalked glands are initiated later in bract growth and undergo development and maturation on medium, to full sized bracts. Glands are epidermal in origin and derived, with one exception, from a single epidermal initial. The capitate-stalked gland is the exception and is of special interest because it possesses a multicellular stalk secondarily derived from surrounding epidermal and subepidermal cells. Glands differentiate early in development into an upper secretory portion and a subtending auxiliary portion. The secretory portion, depending on gland type, may range from a few cells to a large, flattened multicellular disc of secretory cells. The secretory portion produces a membrane-bound resinous product which caps the secretory cells. Capitate-stalked glands are considered to be of particular evolutionary significance because they may represent a gland type secondarily derived from existing capitate-sessile glands.     

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.     

Anatomical and histochemical features of the digestive system of Octopus vulgaris Cuvier, 1797 with a special focus on secretory cells

This study reports a detailed anatomical and histological study of the digestive system of Octopus vulgaris. Emphasis was placed on characterising the glands and glandular cells and their distribution throughout the digestive tract. The use of classic histological and histochemical techniques revealed two morphological types of glandular cells: granular and mucous. Moreover, the histochemical analysis indicated specialisation of mucous glandular cells in the buccal mass, the submandibular gland and the caecum for secreting acid and neutral glycoconjugates. The cells of the anterior salivary glands are specialised for secreting neutral glycoproteins, and those of the posterior salivary glands are specialised for granular and mucous secretion. The oesophagus, crop and stomach lack glandular cells, but both granular and mucous glandular cells are found in the intestine. An unusual structure resembling the typhlosole of bivalves is described for the first time in the intestine of O. vulgaris. The highly ciliated epithelium and location of the structure in the anterior part of the intestine suggest a possible role in bypassing the caecum, stomach and intestine. We discuss how these cells and organs contribute to the process of digestion in the light of the present histological and histochemical data and of previously published information on the morphology and physiology of digestion in the octopus.     

GLAND DISTRIBUTION AND CANNABINOID CONTENT IN CLONES OF CANNABIS SATIVA L.

The relationship between glandular trichomes and cannabinoid content in Cannabis sativa L. was investigated. Three strains of Cannabis, which are annuals, were selected for either a drug, a non-drug, or a fiber trait and then cloned to provide genetically uniform material for analyses over several years. The distribution of the number and type of glands was determined for several organs of different ages including the bract and its subtending monoleaflet leaf and the compound leaf on pistillate plants. Quantitation of glands on these structures was integrated with gas chromatographic analyses of organ cannabinoid profiles. A negative correlation was found between cannabinoid content and gland number for each of the three clones. Isolated heads of the capitate-stalked glands also were analyzed for cannabinoid content and found to vary in relation to clone and gland age. These studies indicate that cannabinoids may occur in plant cells other than glandular trichomes. The results of these studies emphasize the need for stringent sampling procedures in micromorphological studies on trichome distribution and analytical determinations of cannabinoid content in Cannabis.     

杜鹃花属映山红亚属植物叶表皮显微形态特征及其分类学意义

为澄清映山红亚属(Rhododendron subg. Tsutsusi)内的系统关系问题,该研究选择杜鹃花属映山红亚属37种植物作为对象,其中29种为首次报道,通过扫描电镜观察其叶表皮显微特征。结果表明:(1)气孔器均为无规则型且均在远轴面。(2)根据叶片微形态特征将映山红亚属的种类分为杜鹃型(R. simisii-type)、岭南杜鹃型(R. mariae-type)、皋月杜鹃型(R. indicum-type)、崖壁杜鹃型(R. saxatile-type)及丁香杜鹃型(R. farrerae-type)5种类型。(3)杜鹃型植物的气孔器周边无或具间断的条形突起。(4)岭南杜鹃型植物的气孔器周围有多层条形突起环绕,保卫细胞两极不具T型加厚。(5)皋月杜鹃型植物的叶表皮保卫细胞两极具有T型加厚,与叶状苞亚属(Subg. Therorhodion)叶状苞杜鹃(R. redowskianum)的气孔器特征相似,推测其与叶状苞亚属具有一定的亲缘关系。(6)崖壁杜鹃型植物的叶表皮毛单一,多呈卷曲状,与其他类型有所不同且未见腺体。(7)丁香杜鹃型植物的叶表皮仅有腺体。(8)依据叶表皮微形态特征讨论了一些近缘种类的关系,如倾向于将背绒杜鹃(R. hypoblematosum)和千针叶杜鹃(R. polyraphidoideum)处理为独立的种,支持保留紫薇春(R. naamkwanense var. cryptonerve)作为南昆杜鹃(R. naamkwanense)的变种地位,不支持将腺花杜鹃(R. adenanthum)并入细瘦杜鹃(R. tenue)作异名处理等。该研究结果表明叶表皮显微特征在映山红亚属内物种划分上具有重要价值。     

Structure and development of bracteal nectary glands in Aphelandra (Acanthaceae)

A survey of bracteal (extrafloral) nectaries in species of Aphelandra (Acanthaceae) reveals substantial diversity. Each bracteal nectary is an aggregate of individual glands that vary in number, size, and structure among species. Glands contain three cell layers: a palisade-like secretory cell layer, a one-to-many-celled intermediate layer with thickened cell walls, and a foot layer. Members of the A. pulcherrima complex have one of two distinct gland types: relatively small glands with a single-celled intermediate layer or larger glands that have a multicellular intermediate layer. Nectaries composed of small glands are patches of many (>50) glands, whereas those composed of large glands are patches of < 10 glands. Four outgroup species have bracteal nectaries of numerous small glands with pluricellular intermediate layers. Glands of all three types are initiated as single enlarged protodermal cells, and all undergo similar early periclinal divisions; the large-gland type shows greater subsequent enlargement with many more anticlinal divisions. The bracteal nectar glands are interpreted to be homologous with simpler glandular trichomes, and mark a monophyletic lineage within Aphelandra. Comparisons with outgroup species show that both nectary types in the A. pulcherrima complex have diverged from an ancestral condition of numerous small glands with pluricellular intermediate layers. Use of the ontogenetic criterion to polarize gland type within the A. pulcherrima complex would yield erroneous results because evolution has apparently involved a developmental truncation with loss of cell divisions in the intermediate layer of small glands. Comparable nectar glands in more distant taxa are interpreted as remarkable cases of convergent evolution, perhaps from similar trichome precursors.     

The exocrine glands of the male of Eldana saccharina walker (Lepidoptera,galleriinae): Intervention in precopulatory behavior

The scent apparatus of male Eldana saccharina is a glandular complex on the costal area of the forewing. It consists of two parts; glandular complex 1 is composed of five kinds of cells (epidermal cells, scale cells, glandular cells, supporting cells, duct cells); glandular complex 2 also shows five types of cells (epidermal cells, scale cells, glandular cells, duct cells, trichogen cells). The secretory products of the two parts are discharged into separate ducts which converge before opening onto the lower side of the wing. The male also has two prominent hair-pencils borne on the coremata and large secretory trichogen cells on the genital valves. Each of these exocrine gland components plays an important part in formation of the chemically complex pheromones utilized in the precopulatory behavior of the male.     

Ultrastructure of the foregut and associated glands in the lung fluke,Paragonimus miyazakii (Digenea: Troglotrematidae), with particular reference to their functional roles

The foregut and associated glands of a digenetic trematode, Paragonimus miyazakii, were examined in the forebody by transmission and scanning electron microscopy as well as by light microscopy, and their functional roles were discussed. The foregut is lined with a general tegument without spines and sensory receptors throughout its length, although it consists of the mouth, pharynx, and esophagus. This foregut tegument is regionally and intraregionally modified in appearance, suggesting the performance of auxiliary functions in digestion. This appearance is characterized by long, frequent cytoplasmic extensions of the apical tegument around the middle portion of the mouth and the anterior esophagus. Electron-dense granules and multimembranous and multilamellar bodies are developed in the tegument to various degrees, and elaborately in the apical layer of the prepharynx. A single type of unicellular gland is embedded in the antero-middle part of the worm in small groups. The gland cells synthesize clear secretory granules as a chief product, each granule with a pleomorphic, dense, core-like inclusion. Mature granules are elliptical in shape, approximately 500 nm in diameter, and are subsequently discharged into the prepharyngeal foregut lumen after passing through the elongated cytoplasm of the gland cell. In the prepharynx and pharynx, host blood cells are apparently processed for digestion. In the wide lumen of the esophagus, foodstuff could undergo sufficient digestion prior to absorption by the cecal epithelium. J. Morphol. 237:43–52, 1998. © 1998 Wiley-Liss, Inc.     

Biosynthesis of formic acid by the poison glands of formicine ants

The biosynthesis of formic acid in the poison glands of formicine ants is closely related to the C-1 metabolism of the glandular cells. Experiments utilizing radiolabeled amino acids revealed that serine is a major precursor, contributing both its α and β carbons to formic acids. 5,10[¹⁴C]methylene H₄folate and 5,10[¹⁴C]methenyl H₄folate also serve as precursors of formic acid in the poison gland, suggesting that they are intermediates in the pathway. Furthermore, these H₄folate derivatives were isolated from poison glands following incubation with [3-¹⁴C]serine and proved radioactive. The glandular cells are also exceptionally rich in the enzymes responsible for these reactions, supporting the proposed pathway.Although this pathway has been established in various organism, the uniqueness of the poison gland system is that it accumulates formic acid to large extent, yet avoids its cytotoxicity. This is made possible by a combination of the biochemical characteristics of the pathway and the special morphological features of the poison gland.     

MORPHOLOGY OF GLANDULAR HAIRS OF CANNABIS SATIVA FROM SCANNING ELECTRON MICROSCOPY

Three distinct types of glandular hairs of increasing morphological complexity which occur on flowering tops of Cannabis sativa L. (marihuana) are described from scanning electron microscopy. These gland types—termed bulbous, capitate-sessile, and capitate-stalked, described from pistillate plants—occur in greatest abundance on the outer surface of bracts ensheathing the ovary. Bulbous and capitate-sessile glands, which arise at an early stage in bract development, are scattered over the bract surface. Mature bulbous glands have a small swollen head on a short stalk, whereas capitate-sessile glands have a large globular head attached directly to the bract surface. Because of their numbers and large size, capitate-sessile glands are the most conspicuous gland type during the early phase of bract development. Capitate-stalked glands, which have a large globular head on a tall, multicellular stalk, differentiate during subsequent bract development. These stalked glands arise first along the bracteal veins and then over the entire bract surface. A voluminous, fluid secretory product accumulates in the glandular head of all three types. These glands are believed to be a primary site of localization of the marihuana hallucinogen, tetrahydrocannabinol.     

Histochemical evidence of β‐chitin in parapodial glandular organs and tubes of Spiophanes (Annelida,Sedentaria: Spionidae), and first studies on selected Annelida

A generic character of the genus Spiophanes (Annelida, Sedentaria: Spionidae) is the presence of parapodial glandular organs. Parapodial glandular organs in Spiophanes species include secretory cells with cup‐shaped microvilli, similar to those present in deep‐sea inhabiting vestimentiferans and frenulate Siboglinidae. These cells are supposed to secrete β‐chitin for tube‐building. In this study, transverse histological and/or ultrathin sections of parapodial glandular organs and tubes of Spiophanes spp. as well as of Glandulospio orestes (Spionidae) and Owenia fusiformis (Oweniidae) were examined. Fluorescent markers together with confocal laser scanning microscopy, and Raman spectroscopy were used to detect chitin in the parapodial glandular organs of Spiophanes and/or in the glands of Owenia and Glandulospio. Tubes of these taxa were tested for chitin to elucidate the use of it for tube‐building. The examinations revealed a distinct labelling of the gland contents. Raman spectroscopy documented the presence of β‐chitin in both gland types of Spiophanes. The tubes of Spiophanes were found to have a grid‐like structure that seems to be built with this β‐chitin. Tests of tubes of Dipolydora quadrilobata (Spionidae) for chitin were negative. However, the results of our study provide strong evidence that Spiophanes species, O. fusiformis and probably also G. orestes produce chitin and supposedly use it for tube‐building. This implies that the production of chitin and its use as a constituent part of tube‐building is more widespread among polychaetes as yet known. The histochemical data presented in this study support previous assumptions inferring homology of parapodial glandular organs of Spionidae and Siboglinidae based on ultrastructure. Furthermore, transmission electron microscopy‐based evidence of secretory cells with nail‐headed microvilli in O. fusiformis suggests homology of parapodial grandular organs across annelids including Sipuncula. J. Morphol. 276:1433–1447, 2015. © 2015 Wiley Periodicals, Inc.     

The structure of the cutaneous pedal glands in the banded snail Cepaea hortensis (Müller, 1774)

Although gastropods have been crawling through the ocean and on the land for 60 million years, we still know very little about the sticky mucus produced in their foot. Most research has been focused on marine species in particular and, to a lesser extent, on the well‐known terrestrial species Arion vulgaris and Cornu aspersum. Within this study, we aim to characterize the foot anatomy of a smaller representative of the family Helicidae, the banded snail Cepaea hortensis. We are particularly interested in the microanatomy of the foot glands, their position, and the histochemical nature of their secretory content. Characterization of the dorsal foot region of Cepaea hortensis reveals four glands, differing in their size and in the granules produced. Histochemically, three of them react positively for sugars (PAS staining and lectin affinity tests for mannose, glucose and N‐acetyl‐d ‐glucosamine) and acidic proteins (positive Alcian blue and Toluidine blue staining), indicating the presence of acidic glycosaminoglycans. The fourth gland type does not react to any of these dyes. The ventral pedal region includes two different gland types, which are positive for the presence of acidic glycoproteins, with a lectin affinity for mannose only. A comparison with Helix pomatia indicates differences regarding the number of glands and their contents. In Helix, only three gland types are described in the dorsal region of the foot, which show a similar granular appearance but nevertheless differ in their chemical composition. Congruently, there are two gland types in the ventral region in both species, whereas in Helix an additional sugar moiety is found. This raises the question whether these differences between the pedal glandular systems of both helicid species are the result of protection or size‐related adaptations, as they occur in the same habitat.