T.V. Kuzmina

The periesophageal celom of the articulate brachiopod Hemithyris psittacea (Rhynchonelliformea, Brachiopoda)

The celomic system of the articulate brachiopod Hemithyris psittacea is composed of the perivisceral cavity, the canal system of the lophophore, and the periesophageal celom. We study the microscopic anatomy and ultrastructure of the periesophageal celom using scanning and transmission electron microscopy. The periesophageal celom surrounds the esophagus, is isolated from the perivisceral cavity, and is divided by septa. The lining of the periesophageal celom includes two types of cells, epithelial cells and myoepithelial cells, both are monociliary. Some epithelial cells have long processes extending along the basal lamina, suggesting that these cells might function as podocytes. The myoepithelial cells have basal myofilaments and may be overlapped by the apical processes of the adjacent epithelial cells. The periesophageal celom forms protrusions that penetrate the extracellular matrix (ECM) of the body wall above the mouth and the ECM that surrounds the esophagus. The canals of the esophageal ECM form a complicated system. The celomic lining of the external circumferential canals consists of the epithelial cells and the podocyte‐like cells. The deepest canals lack a lumen; they are filled with the muscle cells surrounded by basal lamina. These branched canals might perform dual functions. First, they increase the surface area and might therefore facilitate ultrafiltration through the podocyte‐like cells. Second, the deepest canals form the thickened muscle wall of the esophagus and could be necessary for antiperistalsis of the gut. J. Morphol., 2011.

Structure of the brachiopod lophophore

Data on the development, structure, and functional morphology of the brachiopod lophophore are analyzed. The common origin of the tentacle apparatus in Lophophorata from the postoral ciliary band of the larva is shown. The brachiopod lophophore is based on the brachial axis consisting of the brachial fold running along the row of tentacles. The brachial axis may be attached to the brachial (dorsal) mantle lobe (trocholophe, schizolophe, and ptycholophe lophophores) or extend freely into the mantle cavity to form coiling brachia (spirolophe, zygolophe, and plectolophe lophophores). The circulation of water flows through the mantle cavity in the brachiopods with attached and free lophophores is described. A new hypothesis on the sorting of particles suspended in water during filtration is proposed.

The accessory hearts of the articulate brachiopod Hemithyris psittacea

Abstract Scientists have long known that the articulate brachiopods have accessory hearts in addition to the main heart. However, the critical factors that determine the location of the accessory hearts and their fine structure remained unclear. We studied the location and fine structure of the accessory hearts in the articulate brachiopod Hemithyris psittacea that is necessary for a better understanding of their function. We observed regular arrangement of the accessory hearts behind the nephridial funnels, close to the gonad vessels, but each examined specimen showed an individual pattern of the accessory heart location. The accessory heart wall is composed of outer coelomic epithelium, extracellular matrix, and inner amoebocytes. The coelomic epithelium consists of epithelio-muscle cells with the basal muscle processes, which give rise to the secondary thin non-muscular processes, interdigitating with each other and forming irregular ultrafiltration slits. The inner amoebocytes do not form a continuous layer and lack desmosomes; thus, they cannot be considered as a true epithelium. Brachiopod accessory hearts apparently fulfill two functions. The well-developed muscular net in the accessory heart wall performs the propulsion of blood to branched gonad vessels. The epithelio-muscle cells in the accessory heart wall are the podocyte-like cells; therefore, the accessory heart wall can be interpreted as the ultrafiltration site. The association of accessory hearts with nephridia suggests their involvement in excretion.

The first data on the innervation of the lophophore in the rhynchonelliform brachiopod Hemithiris psittacea : what is the ground pattern of the lophophore in lophophorates?

BackgroundThe nervous system in brachiopods has seldom been studied with modern methods. An understanding of lophophore innervation in adult brachiopods is useful for comparing the innervation of the same lophophore type among different brachiopods and can also help answer questions about the monophyly of the lophophorates. Although some brachiopods are studied with modern methods, rhynchonelliform brachiopods still require investigation. The current study used transmission electron microscopy, immunocytochemistry, and confocal laser scanning microscopy to investigate the nerve system of the lophophore and tentacles in the rhynchonelliform Hemithiris psittacea.ResultsFour longitudinal nerves pass along each brachium of the lophophore: the main, accessory, second accessory, and lower. The main brachial nerve extends at the base of the dorsal side of the brachial fold and gives rise to the cross nerves, passing through the extracellular matrix to the tentacles. Cross nerves skirt the accessory brachial nerve, branch, and penetrate into adjacent outer and inner tentacles, where they are referred to as the frontal tentacular nerves. The second accessory nerve passes along the base of the inner tentacles. This nerve consists of Ʊ-like parts, which repetitively skirt the frontal and lateral sides of the inner tentacle and the frontal sides of the outer tentacles. The second accessory nerve gives rise to the latero-frontal nerves of the inner and outer tentacles. The abfrontal nerves of the inner tentacles also originate from the second accessory nerve, whereas the abfrontal nerves of the outer tentacles originate from the lower brachial nerve. The lower brachial nerve extends along the outer side of the lophophore brachia and gives rise to the intertentacular nerves, which form a T-like branch and penetrate the adjacent outer tentacles where they are referred to as abfrontal nerves. The paired outer radial nerves start from the lower brachial nerve, extend into the second accessory nerve, and give rise to the lateroabfrontal tentacular nerves of the outer tentacles.ConclusionsThe innervation of the lophophore in the rhynchonelliform Hemithiris psittacea differs from that in the inarticulate Lingula anatina in several ways. The accessory brachial nerve does not participate in the innervation of the tentacles in H. psittacea as it does in L. anatina. The second accessory nerve is present in H. psittacea but not in L. anatina. There are six tentacular nerves in the outer tentacles of H. psittacea but only four in all other brachiopods studied to date. The reduced contribution of the accessory brachial nerve to tentacle innervation may reflect the general pattern of reduction of the inner lophophoral nerve in both phoronids and brachiopods. Bryozoan lophophores, in contrast, have a weakened outer nerve and a strengthened inner nerve. Our results suggest that the ancestral lophophore of all lophophorates had a simple shape but many nerve elements.

Larval development of Brachiopod Coptothyris grayi (Davidson, 1852) (Brachiopoda, Rhynchonelliformea)

The larval development of the Brachiopod Coptothyris grayi (Davidson, 1852) from the Sea of Japan is described for the first time. Ciliated blastula proved to represent the first free-swimming stage. The blastopore is initially formed as a rounded hole stretching later along the anteroposterior axis. The larva is first divided into two lobes (the apical lobe and the trunk); the mantle lobe is formed later as two lateral folds. Two pairs of seta bundles appear in the late stage larvae. The apical larval lobe in brachiopods is supposed to match the pre-oral lobe and anterior part of the trunk with tentacles in phoronids.

Ultrastructure of the lophophoral coelomic lining in the brachiopod Hemithiris psittacea: functional and evolutionary significance

The ultrastructure of the lophophoral coelomic lining in the articulate brachiopod Hemithiris psittacea was studied using electron microscopy and confocal laser scanning microscopy. The coelomic system of the lophophore consists of large and small canals; both extend along each brachium. A small coelomic canal gives rise to a blind coelomic canal into each tentacle. The lophophoral coelothelium consists of two types of cells, epithelio-muscle and peritoneal cells, and exhibits different types of organization that is known in the Bilateria: from an epithelium consisting only of epithelio-muscle cells to a pseudostratified myoepithelium. The lophophoral coelothelium forms the musculature and blood vessel wall of the brachia and tentacles. The lophophoral blood vessel runs in the extracellular matrix of the septum that separates the large and small canals. In the lophophoral vessel, the upper wall consists of epithelio-muscle cells, whereas the lower wall is formed by peritoneal cells. The lophophoral vessel gives rise to blind branches into each tentacle. The lophophoral wall is highly muscular and contains both longitudinal and transverse muscles. Because the spirolophe lophophore of H. psittacea is supported only by short crura, the coelomic system of the lophophore is assumed to stabilize the lophophore by regulating the hydrostatic pressure of the coelomic fluid.

Spermatogenesis in the deep-sea brachiopod Pelagodiscus atlanticus and the phylogenetic significance of spermatozoon structure

Details of spermatogenesis and sperm organization are often useful for reconstructing the phylogeny of closely related groups of invertebrates. Development in general and gametogenesis in particular usually differ in shallow water and deep‐sea invertebrates. Here, the spermatogenesis and ultrastructure of sperm were studied in the deep‐sea brachiopod Pelagodiscus atlanticus. The testes of P. atlanticus are voluminous sacs located along the lateral sides of the body. Germ cells develop around the blood capillaries, contact the basal lamina, and contain germ plasm, numerous mitochondria, Golgi apparatus, lipid droplets, and centrioles of the rudimentary cilium. During spermatogenesis, several proacrosomal vesicles appear at the posterior pole of the cell; these vesicles then fuse and migrate to the anterior pole. The spermatozoon has a head with an acrosome, nucleus, eight mitochondria, proximal and distal centrioles orthogonally arranged, and a long tail. Comparative analysis suggests that the spermatozoon of P. atlanticus can be considered the most ancestral among all brachiopods. Such an organization indicates that fertilization is external in this deep‐sea species. Spermatozoa of other brachiopods should be regarded as derived from this ancestral type. The transformation of brachiopod spermatozoa might have occurred in three different ways that correspond to the three main clades of recent brachiopods: Linguliformea, Craniiformea, and Rhynchonelliformea.