Histology and regeneration of the radula of Pomacea bridgesi (Gastropoda,Prosobranchia)

Summary Histology, physiological regeneration, and degradation of the taenioglossan prosobranch radula and its concomitant epithelia were studied by light and electron microscopy (TEM, SEM), electron microprobe analysis, and autoradiography. Taenioglossa have seven multicellular odontoblastic cushions which produce the tooth matrix by apocrine secretion; many long microvilli are also incorporated. In contrast to pulmonates, the odontoblasts of prosobranchs are capable of division, and their mitoses contribute to the expansion of the cushions, but presumably also to the displacement of degenerating odontoblasts. The seven cushions are isolated from each other by separation cells. The radular membrane is produced from microvilli of membranoblasts and a substance secreted at the base of microvilli.Strands of the supraradular epithelium subsequently move in between the teeth and finally enclose them completely. They effect the hardening and mineralization of the teeth. The strands move together with the radula towards the anterior and are extruded at the opening of the radular sheath; their degeneration, however, has already started in the middle section of the sheath. Epithelial cells are produced by two completely separated mitotic centres which lie dorsolaterally at the end of the sheath.In the subradular epithelium, mitotic activity is scattered over the posterior half of the sheath but is not found in the region where the supramedian radula tensor muscle is inserted. The epithelial cells move forward, but at a much lower rate than the radula. At the opening of the sheath the subradular membrane is generated, while cells of the subradular epithelium lying between the lamellae of the subradular membrane are extruded.The subradular membrane is limited to the functional part of the radula. It is situated on the distal radular epithelium, which is obviously not a continuation of the subradular epithelium. In test animals treated with tritiated thymidine, there is a very strong stationary centre of labeled cells at the beginning of the epithelium, but so far no mitoses have been found in this centre and the labeled cells do not move on continually. In the middle of the distal epithelium mitoses do occur, and the labeled cells permit the assumption that these cells do not migrate at all to the anterior end. At least in Prosobranchia, the distal radular epithelium does not transport the radula to its degradation zone. The transport mechanism for the radula is still unknown.     

Degradation of the radula in the snails Biomphalaria glabrata say and Limnaea stagnalis L. (Gastropoda,Pulmonata)

Summary The radula of snails is formed at the posterior end of the radular gland or pocket, and degraded at the same rate at its anterior end. Degradation is due to different secretory activities of the inferior epithelium of the radular gland. Its secretions seem to degrade enzymatically the matrix of the radular membrane and basal plates of teeth, leaving only chitin containing microfibres and degradation products. The sclerotized parts of the teeth remain unchanged, but as they are now only loosely connected with the radular membrane. they are torn off easily during feeding movements. The rest of the degraded and frayed radular membrane and the subradular membrane are also lost by abrasion during feeding. The cells of the inferior epithelium are connected with each other by septate desmosomes and an elaborate system of deep lateral interdigitation which may provide tensile strength. Extrusion of degraded cells of the inferior epithelium into the subradular membrane takes place, although the thick basal lamina forms a continuous sheath which is closely adjoined to the basal parts of the inferior epithelium. Nerve fibres containing vesicles with electron dense neurosecretory material (deduced from the diameter of 200–250 nm) are attached to this sheath or penetrate into it; they may be involved in the regulation of production and degradation processes during radula replacement. Problems of the forward transport of radula and inferior epithelium are discussed.     

Radular mechanosensory neuron in the buccal ganglia of the terrestrial slug,Incilaria fruhstorferi

A radular mechanosensory neuron, RM, was identified in the buccal ganglia of Incilaria fruhstorferi. Fine neurites ramified bilaterally in the buccal ganglia, and main neurites entered the subradular epithelium via buccal nerve 3 (n3). When the radula was distorted by bending, RM produced an afferent spike which was preceded by an axonic spike recorded at n3. The response of RM to radular distortion was observed even in the absence of Ca²⁺, which drastically suppressed chemical synaptic interactions. Therefore, RM was concluded to be a primary radular mechanoreceptor.During rhythmic buccal motor activity induced by food or electrical stimulation of the cerebrobuccal connective, RM received excitatory input during the radular retraction phase. In the isolated buccal ganglia connected to the radula via n3s, the afferent spike, which had been evoked by electrical stimulation of the subradular epithelium, was broadened with the phasic excitatory input. Since the afferent spike was also broadened by current injection into the soma, depolarization due to the phasic input may have produced the spike broadening.Spike broadening was also observed during repetitive firing evoked by current injection. The amplitude of the excitatory postsynaptic potential in a follower neuron increased depending on the spike broadening of RM.Abbreviations CBC cerebrobuccal connective - EPSP excitatory postsynaptic potential - n1,n3 buccal nerves 1 and 3 - RBMA rhythmic buccal motor activity - RM radular mechanosensory neuron - SMT supramedian radular tensor neuron     

Comparative buccal anatomy in Helisoma (mollusca,pulmonata, basommatophora)

Dissections were performed to document buccal anatomy in three species of the pulmonate genus Helisoma Swainson, 1840. The 28 muscles which are responsible for radular feeding in these animals are organized in three concentric and integrated envelopes. The deepest of these includes muscles which manipulate the radula about the odontophoral cartilage. Elements of the middle envelope direct movements of the cartilage within the buccal cavity, and muscles of the outer envelope control movements of the buccal mass within the cephalic haemocoel. Motion analysis by videomicrography showed that muscles of the middle and outer envelopes contribute to the action of radular feeding by acting as antagonists to other muscles and to hydrostatic elements of the buccal apparatus. Observations of radular dentition showed that although each of the three species examined has a unique radula, especially with regard to the specific details of tooth shape, all resemble a radula characteristic of the Planorbidae with regard to other, more general, aspects of ribbon architecture.     

Main patterns of radula formation and ontogeny in Gastropoda

Gastropoda is morphologically highly variable and broadly distributed group of mollusks. Due to the high morphological and functional diversity of the feeding apparatus gastropods follow a broad range of feeding strategies: from detritivory to highly specialized predation. The feeding apparatus includes the buccal armaments: jaw(s) and radula. The radula comprises a chitinous ribbon with teeth arranged in transverse and longitudinal rows. A unique characteristic of the radula is its continuous renewal during the entire life of a mollusk. The teeth and the membrane are continuously synthesized in the blind end of the radular sac and are shifted forward to the working zone, while the teeth harden and are mineralized on the way. Despite the similarity of the general mechanism of the radula formation in gastropods, some phylogenetically determined features can be identified in different phylogenetic lineages. These mainly concern shape, size, and number of the odontoblasts forming a single tooth. The radular morphology depends on the shape of the formation zone and the morphology of the subradular epithelium. The radula first appears at the pre- and posttorsional veliger stages as an invagination of the buccal epithelium of the larval anterior gut. The larval radular sac is lined with uniform undifferentiated cells. Each major phylogenetic lineage is characterized by a specific larval radula type. Thus, the docoglossan radula of Patellogastropoda is characterized by initially three and then five teeth in a transverse row. The larval rhipidoglossan radula has seven teeth in a row with differentiation into central, lateral, and marginal teeth and later is transformed into the adult radula morphology by the addition of lateral and especially marginal teeth. The taenioglossan radula of Caenogastropoda is nearly immediately formed in adult configuration with seven teeth in a row.     

Radula formation in two species of Conoidea (Gastropoda)

The radula is the basic feeding structure in gastropod molluscs and exhibits great morphological diversity that reflects the exceptional anatomical and ecological diversity occurring in these animals. This uniquely molluscan structure is formed in the blind end of the radular sac by specialized cells (membranoblasts and odontoblasts). Secretion type, and the number and shape of the odontoblasts that form each tooth characterize the mode of radula formation. These characteristics vary in different groups of gastropods. Elucidation of this diversity is key to identifying the main patterns of radula formation in Gastropoda. Of particular interest would be a phylogenetically closely related group that is characterized by high variability of the radula. One such group is the large monophyletic superfamily Conoidea, the radula of which is highly variable and may consist of the radular membrane with five teeth per row, or the radular membrane with only two or three teeth per row, or even just two harpoon-like teeth per row without a radular membrane. We studied the radulae of two species of Conoidea (Clavus maestratii Kilburn, Fedosov & Kantor, 2014 [Drilliidae] and, Lophiotoma acuta (Perry, 1811) [Turridae]) using light and electron microscopy. Based on these data and previous studies, we identify the general patterns of the radula formation for all Conoidea: the dorsolateral position of two groups of odontoblasts, uniform size, and shape of odontoblasts, folding of the radula in the radular sac regardless of the radula configuration. The morphology of the subradular epithelium is most likely adaptive to the radula type.     

Suctorial feeding in a deep-sea octopus as a means of niche partitioning (Cephalopoda: Octopodidae)

Molecular phylogenetic analyses indicate that two clades of deep-sea octopuses evolved at opposite ends of the earth to become globally sympatric. Coexistence of these overtly similar, but phylogenetically distinct octopuses requires some means of niche partitioning. To investigate details of feeding, the buccal complexes of specimens of each clade, Muusoctopus and Graneledone, were sectioned at 90° to the radular ribbon. The buccal complex of Muusoctopus is the same as reported in Octopus; the radula and its bolsters extend the length of the buccal complex. In Graneledone, however, the radula and its bolsters are restricted to anterior half of the buccal complex. Posterior to the radular sac, a vertically oriented muscle, named here the buccal abductor, extends from the floor of the mouth to the base of the buccal complex. In Muusoctopus, the bolsters extend the radula to bring food into the mouth; the palps propel it to the esophagus. In Graneledone, although the bolsters extend the radula, contraction of the buccal abductor to expand the posterior mouth may be the primary food mover. The negative pressure differential created draws food into the mouth and to the entry to the esophagus. The buccal abductor may permit the ingestion of larger pieces of prey, as gut contents show. Its evolution may represent a key innovation that heightens deep-sea octopus diversity.     

红条毛肤石鳖齿舌主侧齿中铁还原酶分布及与生物矿化的关系

以红条毛肤石鳖Acanthochiton rubrolineatus(Lischke)齿舌为材料,通过切片和酶组织化学技术,在光镜和电镜下对齿舌主侧齿的微结构及高铁还原酶的存在进行观察,从微观角度了解齿舌主侧齿齿尖的矿化机理。结果显示,成熟主侧齿由齿尖和齿基组成。齿尖结构由外至内分为三层,最外层为磁铁矿层,前后齿面磁铁矿层的厚度不等,后齿面约50μm,前齿面约5-10μm。向内依次为棕红色的纤铁矿层,厚约10μm,及略显黄色的有机基质层,有机基质层占据着齿尖内部的大部分结构。高分辨透射电镜下显示磁铁矿由条状四氧化三铁颗粒组成,长约2-3μm,宽约100-150nm。齿舌的矿化是一个连续过程,不同部段处于不同的矿化阶段,齿舌囊上皮细胞沿囊腔分布,并形成齿片。未矿化的新生主侧齿齿尖中存在由有机基质构成的网状结构。随矿化的进行,有机基质内出现矿物颗粒。初始矿化的齿尖外表面有一个细胞微突层,微突的另一端为囊上皮细胞,矿物质经由微突层达齿尖并沉积于有机基质中,齿尖随之矿化并成熟。初始矿化齿尖的外围有大量的三价铁化物颗粒,稍成熟的齿尖外围同时还出现二价铁化物。新生或初始矿化主侧齿齿尖外围的囊上皮细胞中有大量球形类似于铁蛋白聚集体的内容物,直径0.6-0.8μm,球体由膜包围。齿舌囊上皮组织中存在三价高铁还原酶,此酶分布于上皮细胞的膜表面,可能与齿尖表面磁铁矿的生成有一定的关系。       

Modulation of the feeding system by a radular mechanosensory neuron in the terrestrial slug,Incilaria fruhstorferi

We investigated the modulatory role of a radular mechanoreceptor (RM) in the feeding system of Incilaria. RM spiking induced by current injection evoked several cycles of rhythmic buccal motor activity in quiescent preparations, and this effect was also observed in preparations lacking the cerebral ganglia. The evoked rhythmic activity included sequential activation of the inframedian radular tensor, the supramedian radular tensor, and the buccal sphincter muscles in that order.In addition to the generation of rhythmic motor activity, RM spiking enhanced tonic activities in buccal nerve 1 as well as in the cerebrobuccal connective, showing a wide excitatory effect on buccal neurons. The excitatory effect was further examined in the supramedian radular tensor motoneuron. RM spiking evoked biphasic depolarization in the tensor motoneuron consisting of fast excitatory postsynaptic potentials and prolonged depolarization lasting after termination of RM spiking. These depolarizations also occurred in high divalent cation saline, suggesting that they were both monosynaptic.When RM spiking was evoked in the fictive rasp phase during food-induced buccal motor rhythm, the activity of the supramedian radular tensor muscle showed the greatest enhancement of the three muscles tested, while the rate of ongoing rhythmic motor activity showed no increase.Abbreviations CPG central pattern generator - EPSP excitatory postsynaptic potential - RBMA rhythmic buccal motor activity - RM radular mechanosensory neuron - SMT supramedian radular tensor neuron     

Ultrastructure of possible sites of ultrafiltration in some gastropoda,with particular reference to the auricle of the freshwater prosobranch Viviparus viviparus L.

Summary In the Gastropoda pressure-ultrafiltration of the blood is assumed to be the first step in urine formation. The most probable site of ultrafiltration is the wall of the heart. Since in other animal groups ultrafilters are characterized by a special cell type, the podocyte, the hearts of two pulmonates (Lymnaea stagnalis, Biomphalaria glabrata) and of four prosobranchs (Viviparus viviparus, Bithynia tentaculata, Ampullaria gigas, Littorina littorea) were ultrastructurally investigated, in order to establish whether or not podocytes occur in these structures. It appeared that only in the wall of the auricle of Viviparus podocytes are present. They form a layer underneath the epicardium, the epithelium covering the auricle. It is assumed that in Viviparus ultrafiltration proceeds in the auricle. The possible route of the pro-urine is discussed. The location of the ultrafilters in the other species studied remains still unknown.     

Fine morphology of the jaw apparatus of Puncturella noachina (Fissurellidae,Vetigastropoda)

Jaws of various kinds occur in virtually all groups of Mollusca, except for Polyplacophora and Bivalvia. Molluscan jaws are formed by the buccal epithelium and either constitute a single plate, a paired formation or a serial structure. Buccal ectodermal structures in gastropods are rather different. They can be nonrenewable or having final growth, like the hooks in Clione (Gastropoda, Gymnosomata). In this case, they are formed by a single cell. Conversely, they can be renewable during the entire life span and in this case they are formed by a set of cells, like the formation of the radula. The fine structure of the jaws was studied in the gastropod Puncturella noachina. The jaw is situated in the buccal cavity and consists of paired elongated cuticular plates. On the anterior edge of each cuticular plate there are numerous longitudinally oriented rodlets disposed over the entire jaw surface and immersed into a cuticular matrix. The jaw can be divided into four zones situated successively toward the anterior edge: 1) the posterior area: the zone of formation of the thick cuticle covering the entire jaw and forming the electron‐dense outer layer of the jaw plate; 2) the zone of rodlet formation; 3) the zone of rodlet arrangement; and 4) the anterior zone: the free scraping edge of the plate, or the erosion zone. In the general pattern of jaw formation, Puncturella noachina resembles Testudinalia tessulata (Patellogastropoda) studied previously. The basis of the jaw is a cuticular plate formed by the activity of the strongly developed microvillar apparatus of the gnathoepithelium. However, the mechanism of renewal of the jaw anterior part in P. noachina is much more complex as its scraping edge consists not just of a thick cuticular matrix rather than of a system of denticles being the projecting endings of rodlets. J. Morphol. 275:775–787, 2014. © 2014 Wiley Periodicals, Inc.     

HOMOLOGY OF THE PALLIAL AND PULMONARY CAVITY OF GASTROPODS

The development and morphology of the pallial and pulmonarycavities of various gastropods has been investigated using epoxy-resinserial sections. In the veliger larvae of Cellana sandwicensis(Patellogastro-poda), Gibbula adansonii (Vetigastropoda), Modulustectum (Caenogastropoda) and Ovatella myosotis (Pulmonata) theformation of the pallial cavity is nearly identical. After shellformation a shallow dorsal pallial groove develops beneath themantle edge. During the late veliger stage, the ectoderm formsa deep invagination along the bottom of the pallial groove onthe right side of the larva, giving rise to the pallial cavity.In the ellobiid O. myosotis the pallial cavity becomes the lung(=pulmonary cavity), without any major post-metamorphic modification.Thus, the lung of this species is clearly homologous with thepallial cavity of prosobranchs. The lung of pulmonates withveliger development, as well as of fresh water basommatophoransand stylommatophorans, can be shown to be homologous by comparisonof adult morphology. In contrast to previous views, the pulmonatelung should be regarded as truly homologous with the pallialcavity of prosobranchs and opisthobranchs. In the onchidiidpulmonate Onchidium cf. branchiferum, the larval pallial cavityshifts posteriorly after metamorphosis, where it gives riseto a lung and a cloaca. Contrary to previous interpretations,it can be shown that the onchidiid lung is homologous with atleast part of the pallial cavity. Smeagol climoi has only asmall pallial cavity and no separate lung. The previously described‘lung’ is shown to be a gland. The re-evaluationof the development and morphology of the pulmonate lung hasimportant systematic implications: (1) The pulmonary cavitydoes not represent a synapomorphic character of pulmonates.(2) The gymnomorphs cannot be separated from the remaining pulmonatesbased on lung development. (3) The lack of a lung in the smeagolidsmight give reason to reconsider this group's systematic placementwithin the pulmonates.     

Key innovations in Mesozoic ammonoids: the multicuspidate radula and the calcified aptychus

A nearly complete radula with seven elements per row preserved inside of an isolated, bivalved, calcitic lower jaw (= aptychus) of the Late Jurassic ammonite Aspidoceras is described from the Fossillagerstätte Painten (Bavaria, southern Germany). It is the largest known ammonite radula and the first record for the Perisphinctoidea. The multicuspidate tooth elements (ctenodont type of radula) present short cusps. Owing to significant morphological differences between known aptychophoran ammonoid radulae, their possible function is discussed, partly in comparison with modern cephalopod and gastropod radulae. Analogies between the evolution of the pharyngeal jaws of cichlid fishes and the ammonoid buccal apparatus raise the possibility that the evolution of a multicuspidate radula allowed for a functional decoupling of the aptychophoran ammonoid jaw. The radula, therefore, represents a key innovation which allowed for the evolution of the calcified lower jaws in Jurassic and Cretaceous aptychophoran ammonites. Possible triggers for this morphological change during the early Toarcian are discussed. Finally, we hypothesize potential adaptations of ammonoids to different feeding niches based on radular tooth morphologies.     

General morphology and ultrastructure of the radula of Testudinalia testudinalis (O. F. Müller, 1776) (Patellogastropoda,Gastropoda)

The radular morphology of the patellid species Testudinalia testudinalis (O. F. Müller, 1776) from the White Sea was studied using light, electron, and confocal microscopy. The radula is of the docoglossan type with four teeth per row and consisting of six zones. We characterize teeth formation in T. testidinalis as follows: one tooth is formed by numerous and extremely narrow odontoblasts through apocrine secretion; this initially formed tooth consists of numerous vesicles; the synthetic apparatus of the odontoblasts is localized in the apical and central parts of the cells throughout the cytoplasm and is penetrated by microtubules which are involved in the transport of the synthesized products to the apical part of the odontoblast; the newly formed teeth consist of unpolymerized chitin. Mitotic activity is located in the lateral parts of the formation zone. The first four rows contain an irregular arrangement of teeth, but the radular teeth are regularly arranged after the fifth row. The irregularly arranged teeth early on could be a consequence of the asynchronous formation of teeth and the distance between the odontoblasts and the membranoblasts. The morphological data obtained significantly expands our knowledge of the morphological diversity of the radula formation in Gastropoda.     

Ultrastructure of the supporting cells in the chemoreceptor areas of the tentacles of Pomatias elegans (Müller) (mollusca,prosobranchia) and the ommatophore of Helix pomatia L. (Mollusca,Pulmonata)

The ultrastructure of the supporting cells in the chemoreceptor areas of the tentacles of Pomatias elegans and Helix pomatia is very similar. Complex apical structures are present, and the lateral plasma membrane exhibits three zones: (1) a zone of slight interdigitations; (2) a zone characterized by longitudinal plicae; (3) a zone of basal radiculae. The portions of the sensory cells located within the epithelial layer are accommodated in longitudinal grooves in the supporting cells. However, there are also differences. In Pomatias elegans the apical surface is differentiated into long microvilli that are sometimes dichotomously branched and invested by a surface coat along their entire length. Cytofilia and cilia of the sensory cells pass through this layer of microvilli and surface coat throughout its entire width. In Helix pomatia the supporting cells are somewhat smaller and the apical differentiation consists of candelabra-like protrusions, which are usually three times dichotomously branched. The final branchings, corresponding to microvilli, are called terminal twigs. They are covered by a surface coat, which forms a feltwork. The cytofilia and cilia of the sensory cells that intertwine among the protrusions are confined to the space below the terminal twigs, where they compose the spongy layer.     

Expression of the Intermediate Filament Keratin Gene,K15,in the Basal Cell Layers of Epithelia and the Hair Follicle

The intermediate filament keratin, K15, is present in variable abundance in stratified epithelia. In this study we have isolated and characterized the sheepK15gene, focusing on its expression in the follicles of sheep and mice. We show thatK15is expressed throughout the hair cycle in the basal layer of the outer root sheath that envelops the follicle. Strikingly, however, in large medullated wool follicles, a small group of basal outer root sheath cells located in the region thought to contain hair follicle stem cells areK15-negative. In the follicle bulbK15is expressed in cells situated next to the dermal papilla but not in the inner bulb cells. Elsewhere,K15is expressed at a low, variable level in the basal layer of the epidermis and sebaceous gland, often in a punctate pattern. In the esophagus of the sheepK15expression is restricted to the basal layer, in contrast to human esophagus where it is expressed throughout the epithelium. Transgenic mouse lines established with a 15-kb sheepK15gene construct exhibited faithful expression and showed no phenotypic consequences ofK15overexpression. An investigation of transgene expression showed thatK15is continuously expressed in outer root sheath cells during the hair cycle. Given its expression in the mitotically active basal cell layers of diverse epithelia and the follicle,K15expression appears to signal an early stage in the pathway of keratinocyte differentiation that precedes the decision of a cell to become epidermal or hair-like.     

Olfactory metamorphosis in the coastal tailed frog Ascaphus truei (Amphibia,Anura, Leiopelmatidae)

The structure of the olfactory organ in larvae and adults of the basal anuran Ascaphus truei was examined using light micrography, electron micrography, and resin casts of the nasal cavity. The larval olfactory organ consists of nonsensory anterior and posterior nasal tubes connected to a large, main olfactory cavity containing olfactory epithelium; the vomeronasal organ is a ventrolateral diverticulum of this cavity. A small patch of olfactory epithelium (the “epithelial band”) also is present in the preoral buccal cavity, anterolateral to the choana. The main olfactory epithelium and epithelial band have both microvillar and ciliated receptor cells, and both microvillar and ciliated supporting cells. The epithelial band also contains secretory ciliated supporting cells. The vomeronasal epithelium contains only microvillar receptor cells. After metamorphosis, the adult olfactory organ is divided into the three typical anuran olfactory chambers: the principal, middle, and inferior cavities. The anterior part of the principal cavity contains a “larval type” epithelium that has both microvillar and ciliated receptor cells and both microvillar and ciliated supporting cells, whereas the posterior part is lined with an “adult‐type” epithelium that has only ciliated receptor cells and microvillar supporting cells. The middle cavity is nonsensory. The vomeronasal epithelium of the inferior cavity resembles that of larvae but is distinguished by a novel type of microvillar cell. The presence of two distinct types of olfactory epithelium in the principal cavity of adult A. truei is unique among previously described anuran olfactory organs. A comparative review suggests that the anterior olfactory epithelium is homologous with the “recessus olfactorius” of other anurans and with the accessory nasal cavity of pipids and functions to detect water‐borne odorants. J. Morphol. 2011. © 2011 Wiley Periodicals, Inc.     

The ultrastructure of the blood vessels of Branchiostoma lanceolatum (Pallas) (Cephalochordata)

Summary The ultrastructure of the blood vessels of Branchiostoma has been studied using selected characteristic vessels as examples. It is shown that the vessels are a part of the original blastocoelic cavity and are delimited either by the basal laminae of adjacent epithelia or by connective tissue developed in the blastocoelic space. A brief account of the kinds of connective tissue is given. The observed contractility of some vessels depends on two types of contractile filaments situated in the basal part of the surrounding coelomic epithelia. Amoebocytelike cells are present in the blood. They may sometimes lie in contact with the wall of the vessels or with each other, but never form a typical endothelium with junctional complexes and a basal lamina of its own. Actually, there is no endothelium in any part of the vascular system. It is suggested that the term endothelium should be reserved for a closed cellular lining (with junctions) on the luminal side of the vessel wall, standing on a basal lamina of its own and forming a barrier for the exchange between blood and surrounding tissue. It is concluded that the principal structure of the vascular system of Branchiostoma is different from that of vertebrates, but the same as that of other coelomate invertebrates. The blood vessels in these animals are typically delimited directly by a basal lamina secreted by epithelia (epidermal, coelomic or intestinal) lying peripheral to this lamina, and a true endothelium is not present (with a few questionable exceptions).Abbreviations ac atrial cavity - ace atrial epithelium - ao aorta - ap atrial plexus - ax axon bundle - bc blood cell - bl basal lamina - bl ₁ basal lamina of intestinal epithelium - bl ₂ basal lamina of visceral coelomic epithelium - bl ₃ basal lamina of parietal coelomic epithelium - bl ₄ basal lamina of atrial epithelium - bll basement lamella - cf contractile filaments - co coelomic cavity - coe coelomic epithelium - coe _p parietal coelomic epithelium - coe _v visceral coelomic epithelium - ct dense connective tissue - dv longitudinal dorsal vessel - ep epidermis - epe epipharyngeal groove epithelium - epg epipharyngeal groove - fb fibroblast (?) - fi collagen fiber - fl fibril layer - go gonad - hd hemidesmosome - ie intestinal epithelium - in intestine proper - ip intestinal plexus - iv afferent intestinal vessel - ld liver diverticulum - lu vascular lumen - me myocoelic epithelium - ml muscle lamella - mp myoseptal plexus - ms myoseptum - my myomer - myc myocoelic cavity - nc notochord - ns notochordal sheath - ph pharynx - suc subchordal coelom - sv subintestinal vessel - svv segmental ventral vessel - vv longitudinal ventral vessel Supported by a grant from the Danish Natural Science Research Council     

Development of the pedicle in the articulate brachiopod Terebratalia transversa (Brachiopoda,Terebratulida)

Summary The development of the pedicle in the articulate brachiopod Terebratalia transversa has been examined by electron microscopy. The posterior half of the free-swimming larva comprises a non-ciliated pedicle lobe that contains the primordium of the juvenile pedicle at its distal end. During settlement at five to six days post-fertilization, the pedicle lobe secretes a sticky sheet that attaches the larva to the substratum. As metamorphosis proceeds, the epithelium in the posterior half of the pedicle lobe produces a thin overlying cuticle, and the pedicle primordium develops into a stalk-like anchoring organ. The juvenile pedicle protrudes through the gape that occurs between the posterior margins of the shell valves. A cup-like canopy, called the pedicle capsule, lines the posterior end of the shell and surrounds the newly formed pedicle. The core of the juvenile pedicle is filled with a solid mass of connective tissue. Numerous tonofibrils occur in the pedicle epithelium, and the overlying cuticle consists of amorphous material covered by a thin granular fringe. By one year post-metamorphosis, a body cavity develops anterior to the pedicle. Two pairs of adjustor muscles extend from the posterior end of the shell and traverse the cavity to insert in the pedicle. The connective tissue core of the pedicle in sub-adult specimens lacks muscle cells but contains numerous fibroblasts and collagen fibers. Three regions are recognizable in the connective tissue compartment of the adult pedicle: a subepithelial layer of non-fibrous connective tissue, a central fibrous zone, and a proximal mass of tissue that resembles cartilage.List of abbreviations as adhesive sheet - bc body cavity - bv brachial valve of shell - cf collagen fibrils - ct connective tissue - cu cuticle - di diductor muscle - ec epithelial cell - f fibroblast - fz fibrous zone - g gut - gc granular cell - gd gastric diverticulum - ht hinge tooth - ia interarea of pedicle valve - icl inner cuticular layer - lo lophophore - lu lumen of gut - m mesenchyme - ma mantle - ml mantle lobe - ocl outer cuticular layer - p periostracum - pc pedicle capsule - pce pedicle capsule epithelium - pcl pedicle collar of shell - pcn pedicle connectives - pd pedicle - pe pedicle epithelium - pl pedicle lobe - pv pedicle valve of shell - pzc proximal zone of cartilage-like tissue - s substratum - sel subepithelial layer - t tendon - tf tonofibril - vam ventral adjustor muscle