Functional anatomy of the respiratory system of Branchipolynoe species (Polychaeta, Polynoidae), commensal with Bathymodiolus species (Bivalvia, Mytilidae) from deep-sea hydrothermal vents

 The gills of three species of Branchipolynoe have been studied in order to better understand the morphological and anatomical adaptations of their respiratory system. These Polynoidae live commensally inside the pallial cavity of different species of Bathymodiolus (Mytilidae), found clustered near deep-sea hydrothermal vents and cold seeps, and which harbor chemolithoautotrophic bacteria in their gills. As the mussels exploit hydrothermal fluid, the pallial cavity is perfused with a sulfide-rich hydrothermal water. The gills of Branchipolynoe species are well-developed branched outgrows of the body wall, located on the parapodia, and filled with coelomic fluid. They do not contain blood vessels. Living animals are red, due to the presence of extracellular hemoglobins in the coelom. The gill epidermis is made of supporting cells and a few ciliated cells arranged in longitudinal rows along the branches. Myoepithelial and ciliated cells line the interior of the coelomic cavity which contains the respiratory pigments. Coelomic fluid circulation inside the gills and body cavity is probably facilitated by both the cilia and myoepithelial contractions. The cuticle, the epidermis, and the coelomic epithelium are completely devoid of bacteria. The gill surface areas per unit body weight and the minimum diffusion distances, between external milieu and coelomic hemoglobins, have been calculated and compared with data already obtained on vascular gills of littoral or hydrothermal species of Polychaeta. In Branchipolynoe species, the respiratory surface area is very large, similar to that of a free-living hydrothermal species Alvinella pompejana, and the minimum diffusion distance is short, similar to that of the littoral species Arenicola marina. Although the organization of these coelomic gills in Branchipolynoe species is totally different from that of usual vascular gills, their characteristics represent a unique and effective respiratory system in Polynoidae which has adapted to the hypoxic and sulfide-rich micro-habitat which probably holds in the mantle cavity of vent mussels. In the gill epidermis, numerous secondary and large compound lysosomes are present which might be involved in sulfide detoxification. Accepted: 5 August 1998     

Sanitation with sponge and plunger: western Atlantic slit-wormsnails (Mollusca: Caenogastropoda: Siliquariidae)

Siliquariid Tenagodus (synonym: Siliquaria) snails are obligatory sponge commensals, with uncoiling and laterally slitted shells embedded inside Demospongiae hosts, and shell apertures facing the outside allowing for respiratory and gill‐filter‐feeding activities. Live‐collected animals observed in situ within hosts provide the first detailed functional morphological study of the group, and form the basis for systematic revision of the western Atlantic members of type genus Tenagodus (Siliquariidae, synonym: Tenagodidae). Three species are recognized. Overall anatomical features of wide‐ranging Atlantic T. modestus (Dall, 1881) and T. squamatus (de Blainville, 1827) [ = T. anguillae Mörch, 1861] are similar to those previously described for New Zealand Pyxipoma. A third species, Tenagodus barbadensis sp. nov. , is described from Barbados, based on shell characters. The three species live embedded in a few species of halichondriid and thrombid sponges at depths ranging from c. 20 to several hundred metres and show several characters not previously documented for the family. These include a short compressed‐S‐shaped osphradium, an anal opening positioned in the posterior mantle cavity, and cuspidate inner marginal radular teeth. Presence of shell septa sealing earlier parts of the shell, first described in the early 1800s but subsequently disputed, was verified. Observations on living T. squamatus demonstrated the use of the large tight‐fitting operculum in plunger‐like fashion, flushing the mantle cavity and discharging waste material (including faeces from the posteriorly located anus) through the shell slit into the host sponge's aquiferous canal system. Uncoiling shells, shell and mantle slits, and specialized opercular, gill and gut morphology are interpreted as adaptive traits of the sessile mode of life embedded in sponge tissue. Actively maintained linkage between snail and sponge water‐flow systems provides mutual benefits; these include the allowance for rapid defensive withdrawal and return‐to‐feeding position of the snail (by using the sponge canal system as a vent or reservoir for displaced water), sanitation of the mantle cavity (waste material can leave the snail through the shell slit at the posterior end of the mantle cavity, avoiding fouling of the gill‐feeding system), and by providing increased water flow and, potentially, extra nutritional particles for the sponge (food and waste particles stemming from the snail's activity). The lectotypes selected herein are Serpula anguina Linnaeus, 1758, Tenagodus anguillae Mörch, 1861, and Siliquaria modesta Dall, 1881. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 307–333.     

Morphology and ultrastructure of the gills of terrestrial crabs (Crustacea,Gecarcinidae and Grapsidae): Adaptations for air-breathing

Summary The terrestrial crabsGeograpsus grayi, Geograpsus crinipes, Cardisoma hirtipes andGecarcoidea natalis have a reduced number of gills and show a reduced planar gill surface (SA) compared to aquatic species. Gill lamellae are stiffened and thickened (increasing blood/gas (BG) diffusion distances) and nodules maintain wide spacing between lamellae. Haemolymph is directed through the gill lamellae by rows of pillar cells and in the afferent region an intralamellar septum splits the haemolymph into two parallel networks. Gaps in the lines of pillar cells allow movement of haemolymph between adjacent channels. The afferent vessel distributes haemolymph to the lamella via a number of direct channels including the marginal canal and in large gills with the aid of a long, forked sinus which supplies the ventral and central regions of the lamellae. The marginal canal functions in both distribution and collection of haemolymph; the role varies with species. Potential flow-control sites were identified at the junctions between afferent and efferent areas and where the efferent channels enter the efferent branchial vessel. Each gill receives a branch from the sternal artery which supplies all the lamellae. Transport epithelia is the principal cell type in the gills of all species examined though its location varies between species, either being confined to certain gills or specific parts of the lamellae.The gill lamellae of air-breathing crabs are clearly modified to breathe air (stiffening and presence of nodules), though the overall contribution of the gills to gas exchange has been reduced (smaller SA and longer BG diffusion distances). The role of the gills in air-breathing crabs thus appears to have switched from one of an efficient aquatic gas-exchanger (thin with large surface area) and transport tissue, to one that is predominantly set up for ion-regulation.Abbreviations a afferent branchial vessel - ac afferent channels - art arteriole - ass artifactual subcuticular space - bl basal lamina - c cuticle - col collagen - ct connective tissue - e efferent branchial vessel - ec efferent channels - epi epithelium - f folds - g Glycogen - h haemolymph - hc haemocyte - is intralamellar septum - m marginal canal - mi mitochondria - mt microtubules - n nucleus - p pillar cell - s shaft of efferent vessel - sd septate desmosome     

Structural organization of the gills in pipefish (Teleostei,Syngnathidae)

Summary The morphology and structural features of the gills of the two Western Baltic pipefish Nerophis ophidion and Syngnathus rostellatus were investigated using scanning electron microscopy. The general anatomy of the gills complies with the general pattern in fish. Several adaptations though, show the highly specialized nature of pipefish gills. The filaments are extremely short, few in number and carry only a few lamellae due to the limited space in the branchial cavity. The lamellae have a widely projecting form yet still have a small area in comparison to other fish. Gill irrigation is performed by a specialized pumping mechanism which forces respiratory water through the small but densely packed gill sieve. Although both species live in the same habitat and belong to the same family, differences in gill morphology were found and are related to different lifestyles. S. rostellatus is the more active species and therefore has more filaments per gill arch, more lamellae per filament, wider projecting lamellae and a more extreme utilisation of available space in the gill cavity through a very densely packed gill sieve. N. ophidion has a stationary mode of life and therefore has a less extreme gill anatomy.     

Water flow controls distribution and feeding behavior of two co-occurring coral reef fishes: I. Field measurements

The interaction of flowing water with reef topography creates a continuum of flow microhabitats that can alter species distributions directly via transport of organisms or propagules, or indirectly by modulating the availability of critical resources. To examine how water flow affects the distribution and feeding performance of two species of planktivorous tube blennies (Chaenopsidae), flow speed and turbulence were measured within the feeding areas of Acanthemblemaria spinosa and A. aspera at three sites within Glover’s Reef, Belize. Although co-occurring, A. spinosa occupies topographically high locations (e.g., upright coral skeletons) while A. aspera occupies topographically low shelters in the coral pavement. Boundary layer theory predicts that A. spinosa should experience higher flow (and a higher flux of planktonic food) relative to A. aspera; however, complex topography and oscillatory flow require that this prediction is tested directly in the field. Within each site, the flow experienced by A. spinosa was, indeed, faster and more turbulent than that experienced by A. aspera at site-specific intermediate wave heights. When waves were small, gentle velocity gradients produced similar flows for the two species. When waves were high, flow was uniformly fast through the water column due to thinning of the benthic boundary layer. Plankton availability was similar for the species, with the exception of a greater abundance of harpacticoid copepods at the shelters of A. aspera. Quantitative behavioral observations suggest that the foraging strategies employed by the two fishes exploit the prevailing hydrodynamic conditions. For example, A. spinosa, the stronger swimmer of the two, attacks nearly 100% of the time in the water column where it can exploit the higher flux of plankton associated with faster flows, while A. aspera attacks primarily toward the reef surface where currents are likely to be slower and it can exploit more abundant benthic prey. Communicated by Biology Editor Dr. Mark McCormick     

Species of the water-fern megaspore genus Molaspora from a Cenomanian deposit in western France: occurrence,sporoderm ultrastructure and evolutionary relationships

The partially reticulate sculpture of Molaspora aspera sp. nov., a marsileaceous megaspore from a Cenomanian deposit in western France, distinguishes it from other species of Molaspora. An acrolamella entirely surrounds and obscures a small tetrad scar, a feature that has been demonstrated hitherto within members of the genus only in M. fibrosa. It was also encountered for the first time in M. lobata, with which the new species is associated in the same French mesofossil assemblage. The ultrastructure of the sporoderm of M. aspera is similar to that of M. lobata, but differs particularly in that the inner epispore is markedly thicker and may also contain numerous large, homogeneous spherules or, alternatively, holes of comparable dimensions and only a few small spherules. It is possible that these are a response to some hostile bacterial or other activity when the developing sporoderm was partially permeable. The cavity replacing part of the epispore in one of the specimens, and in the specimen of M. lobata examined, may be a preservational feature or have served to increase buoyancy of the spore in water. Molaspora lobata is very similar to megaspores of fossil and extant Regnellidium, but M. aspera bears some resemblance to other members of extant Marsileaceae and certain species of Cretaceous Arcellites, although there are significant differences between them. This suggests that Molaspora is a heterogeneous taxon embracing megaspores produced by water ferns of more than one natural genus, of which only Regnellidium has survived to the present day.     

Chemical nature of the egg shell in helminths. II. Mode of stabilization of egg shells of monogenetic trematodes

The chemical nature of the egg shells of the monogenetic trematodes Pseudomicrocotyle sp. and Pricea multae, parasitizing the gills of the marine fish Scomberomorus guttatus was studied with a view to understanding the mode of stabilization of the egg shells.The egg shells of the monogenetic trematodes appeared to contain dimers of tyrosine; a conclusion based on fluorescence properties, affinity to toluidine blue, methylene blue, and a positive test for aromatic amino acids. In addition, they are stabilized by -S-S- linkages as indicated by the positive reaction to tests for disulfides. This is also suggested by the shells being refractile to stain but which can be reversed after treatment with sodium thioglycollate.Absence of phenolic tanning in the egg shells of the monogenetic trematodes is indicated by the negative reaction to tests for quinones, the failure to develop color on incubation in catechol after heat treatment, and by the fluorescence in uv light.In these respects the egg shells of the monogenetic trematodes strongly resemble the egg shell of Fasciola hepatica.     

Adrenergic innervation of the gills of brown and rainbow trout,Salmo trutta and S. gairdneri

The adrenergic innervation of structures in the gills of brown and rainbow trout was studied with catecholamine fluorescence histochemistry. In the arterio-arterial vascular pathway, there was an innervation of the afferent and efferent lamellar arterioles, but the afferent and efferent filamental arteries and the secondary lamellae were devoid of any fluorescent nerve fibres. In S. trutta only, there was an additional innervation of the afferent and efferent branchial arteries and the base of the efferent filamental artery. The innervation of the arterio-venous vascular pathway was similar in both trout species. Many fluorescent nerve fibres were found on nutritive arterioles in the gill arch and interbranchial septum, and in the core of each filament between the surface epithelium and the wall of the filament venous sinus. No fluorescent nerve fibres were observed at the origins of the capillaries arising from the efferent filamental artery. The sympathetic nerve supply is provided to the gills mainly through the posttrematic nerve, with an occasional small contribution through the pretrematic nerve. The presence of adrenergic nerves in the gills is discussed in relation to the regulation of blood flow through the arterio-arterial and arterio-venous pathways.     

Morphological and chemical characterization of mineral concretions in the freshwater bivalve Anodonta cygnea (Unionidae)

The freshwater mussel Anodonta cygnea is commonly used as a model organism for biomineralization studies, its peculiar morphofunctional properties also make it an excellent environmental biomonitor. The first detailed on the calcareous concretions from gill and mantle tissue, as well as fluids of the freshwater bivalve A. cygnea, supported by histological, scanning, spectrometry, and spectroscopy analyses. Through these analyses, the morphology, structure, and chemical characterization of these biomineral concretions were accomplished. The concretions represent a high percentage of the dry weight of these organisms. In gill tissue, it can reach up to 50% of dry weight prior to reproductive maturity. Analysis of elemental composition of the tissue concretions showed the presence of calcium and phosphate, as main components, associated with other residual elements like iron, manganese, magnesium, and zinc. Concretions are arranged in concentric alternated layers of organic and inorganic matrix. The shape and size of the concretions vary substantially, from very small, less than 1 μm diameter with very regular round structure, found mainly in the mantle tissue, to more than 50 μm length with irregular globular clusters, found predominantly in the gills. The microstructural organization is of a hydroxyapatite polymorphism in the mantle, in contrast to the gills, which exhibit irregular structure and carbonated hydroxyapatite polymorphism. These differences are supported by higher contents of dinitrogen pentoxide, magnesium, and iron in the mantle concretions, but higher contents of manganese and zinc in the gills. Furthermore, the results indicate that the mineral concretion formation in A. cygnea is a hemocytes reaction to particle or toxic invasions. A second relevant role, concerns the close involvement of these microspherules on the adult and larval shell calcification. J. Morphol. 276:65–76, 2015. © 2014 Wiley Periodicals, Inc.     

The morphology of gills of Haliotis tuberculata (Linnaeus, 1758)

Manganaro, M., Laurà, R., Guerrera, M.C., Lanteri, G., Zaccone, D. and Marino, F. 2011. The morphology of gills of Haliotis tuberculata (Linnaeus, 1758). —Acta Zoologica (Stockholm) 93 : 436–443. Although the morphology of abalone gills has been studied by some authors, up to date no data are available about the gills of Haliotis tuberculata. This study was carried out, by light and electron microscopy, on 10 wild adult H. tuberculata. Gills lamellae produce an undulated surface increasing the area in contact with water. At the level of skeletal rods, we observed a joint‐like structure that allows a checked movement. The left ctenidium is always decidedly larger than the right, probably because of the enormous size of the shell muscle. The cilia permit oxygenated water that leaves the afferent border and is thrust away at the tips of the lamellae by the extremely long cilia. Ciliary movement may take part in sweeping mucous secretions to capture extraneous particles and remove them from the gills. Three types of mucous cells are distributed along the epithelium of the afferent and efferent zones of the gill filament. They seem to play a role in the cleansing of gills in coordination with the muscle contraction and ciliary movement. The presence of microvilli on particular cells reflects their role associated with the absorption of substances from the environment. A haemolymphatic vessel is located in the central zone of the gill filament. The backbone of the haemolymphatic vessel is a chitino‐like structure, which gives support to the gills.     

The Ulrastructure of the Gill of the Lugworm Arenicola marina (L.) (Annelida,Polychaeta)

Arenicola marina gills are hollow, branched, body outgrowths with a central coelomic cavity and afferent and efferent vessels. The gill surface area per unit body weight is about 4 cm²/g wet weight. The blood vascular system anatomy differs from the tip to the base of the gill. In the distal branches of the gill the superficial afferent and efferent vessels are joined by connecting vessels. All vessels arise as spacings between the basal laminae of the thin epidermis and of the coelomic myoepithelium. The contractile part of this epithelium mainly borders the afferent and efferent vessels, whereas pedicel-like cytoplasmic processes extend from the cell bodies and mainly line the connecting vessels. In the proximal branches of the gill the afferent and efferent vessels located in the coelomic cavity are surrounded by the coelomic myoepithelium, and a peripheral blood plexus is present below the epidermis. The gill epidermis is everywhere thin and does not exhibit the characters of a transporting epithelium. The gill coelomic myoepithelium has several functions: (i) periodic contractions of the gill, propelling blood and coelomic fluid toward the central vascular and coelomic compartments; (ii) blood ultrafilration toward the coelomic cavity; (iii) probably transport, suggested by the specialized structures of the lateral membranes of the cells.     

Elongation and mat formation of <Emphasis Type="Italic">Chara aspera</Emphasis> under different light and salinity conditions

Former laboratory results indicate that shoot elongation at low light intensities of Chara aspera is absent already at 10 psu which is within the physiologically optimal salinity range for brackish water populations. To investigate if similar restrictions occur in the field, density and morphology of C. aspera were compared between three freshwater and three brackish water sites along its depth range. The lower depth limit of C. aspera varied considerably among sites (30–600 cm) related to turbidity. Light availability at the lower depth limit corresponded to about 15% of surface irradiance in freshwater and brackish water with lower salinity (3.4 psu). Total length increased and fresh weight:length ratio decreased with depth at these sites indicating shoot elongation related to lower light availability. Due to shoot elongation, light availability was far higher at the upper parts of the shoot than at the bottom in the turbid sites. Light availability at the lower depth limit was higher (about 40%) at two sites with higher salinity (7–8 psu), where no shoot elongation was observed at the lower depth limit. Instead, the plants were stunted and often covered with filamentous algae or shaded by other rooted submerged macrophytes indicating competitive disadvantages of C. aspera at higher salinities. As growth in high densities (mat formation) exposes the plants to severe self-shading, it is suggested that shoot elongation is a prerequisite to mat formation. Dense vegetation of C. aspera was found only in freshwater and brackish water with lower salinity. Single, richly branched plants occurred in clearwater sites with higher salinity. C. aspera was not found in “double stress” environments with both high turbidity and high salinity: We asume that the species is a poor competitor under these conditions. Our results indicate that morphological differences between freshwater and brackish water populations of C. aspera are at least partly explained by salinity rather than genetic differences.     

Geographical variation in habitat choice and host suitability in the parasitoidAsobara rufescens

In the Netherlands, Asobara rufescens (Förster) (Hymenoptera: Braconidae) is a parasitoid of drosophilid larvae in decaying plant material. In several places in the Mediterranean, parasitoids looking very similar to A. rufescens were collected on fermenting substrates. A hybridization experiment showed that the parasitoids were indeed A. rufescens. In an olfactometer Portuguese A. rufescens do not have a preference for either the odour of yeast or decaying leaves, while their Dutch conspecifics prefer the odour of decaying leaves. The survival probability of Portuguese A. rufescens in Drosophila melanogaster Meigen (Diptera: Drosophilidae), a species typical for fermenting substrates, is much higher than the survival probability of Dutch A. rufescens in this host species. It is hypothesized that decaying plant material may be unsuitable for drosophilid larvae during part of the year in the Mediterranean, forcing A. rufescens there to broaden its microhabitat choice. The use of fermenting substrates brings A. rufescens in contact with its sibling A. tabida Nees, a species typical for fermenting substrates in most of Europe. Portuguese A. rufescens appear to be genetically isolated from A. tabida. In the Netherlands, where the two species occupy different microhabitats, there is only a premating barrier.     

ORIGINAL ARTICLE: Mass extinctions in the Azores during the last glaciation: fact or myth?

Aim The influence of the last glaciation on the shallow‐water marine malacofauna of the Azores Islands is reviewed. We test, for this fauna, the ‘Pleistocene temperature theory’ of J.C. Briggs, which hypothesizes that a (supposed) lack of endemics in the older (Azorean endemic) fauna resulted from extinctions caused by a severe drop in sea surface temperatures during the Pleistocene. Location Santa Maria Island, Azores, Portugal. Methods We compare the fossil mollusc fauna of Prainha, Praia do Calhau and Lagoinhas Pleistocene outcrops with the recent mollusc fauna of the Azores Islands. We dated the fossil fauna, using shells of Patella aspera Röding, 1798, by standard U/Th methodology at the GEOTOP laboratory (Université du Québec à Montreal, Canada). Results Dating of the shells of P. aspera indicates that the deposition of the lower unit of the Prainha outcrop corresponded to Marine Oxygen Isotope Substage 5e (MISS 5e). Not a single endemic Azorean species of mollusc that is present in the Pleistocene fossil record has since become extinct, and we found no signs of ‘mass extinctions’ in the littoral marine molluscs of the Azores. The only species that were extirpated from these islands were thermophilic molluscs and littoral bivalves living in fine sand. Main conclusions Our results do not support Briggs’‘Pleistocene temperature theory’. Nor did we find evidence supporting the hypothesis that most of the marine organisms now present in the Azores recolonized the islands after the last glacial maximum.     

Sex change and spatial distribution pattern in an intertidal holothurian Polycheira rufescens in the reproductive season

The patterns of sex change and spatial distribution in an intertidal holothurian, Polycheira rufescens (Chiridotidae; Echinodermata), were investigated on a stony beach in Amakusa, western Kyushu. Field caging experiments revealed that some individuals of P. rufescens underwent a sequential sex change from male to female via hermaphrodite stages and back to male again within a single reproductive season. The sex ratio of the population gradually changed from male dominance at first to equal proportions of males and females as the reproductive season progressed. Toward the end of the reproductive season, immature or spent individuals increased in proportion. Stone size appeared to be an important factor affecting the occurrence of P. rufescens individuals on a stony beach. Analysis of spatial distribution by means of Morisita's index of dispersion and nearest neighbor distances indicated that (i) males showed a slightly stronger tendency to aggregate than females, while females had a tendency toward uniform distribution; (ii) females tended to attract males, as shown by relatively short female-to-male distances; and (iii) there was a substantial variation in male-to-female distances, such that some males were positioned close to females while others were not. It may be suggested that spatial distribution of P. rufescens individuals during the reproductive period is partly dictated by the differential needs of individuals of different sexual states. Received: January 25, 1999 / Accepted: June 10, 1999     

A morphological study on gills of the brown shrimp, Penaeus aztecus

The gills of Penaeus aztecus were examined by light and electron microscopy. They are dendrobranchiate, consisting of a central axis with biserially arranged branches that subdivide into bifurcating filaments. A septum divides the lumina of these structures into afferent and efferent channels. Hemolymph from the sternal sinus flows through the afferent channels into the filaments where it is directed into the efferent channels and finally to the pericardial cavity. In addition to these channels, numerous blood vessels permeate the gill. The cuticle covering the gill overlies a thin epithelium which is separated from hemolymph by a basal lamina. The epithelium, which is active in cuticle secretion, has a series of pillar processes that form subcuticular lacunae. The apical membranes of epithelial cells become folded in shrimp exposed to hypo- and hyperosmotic salinities. Granular cells that contain elaborate Golgi apparati and several types of granules are present throughout the gill. Nephrocytes resembling glomerular podocytes line the efferent channels. A large nerve traverses the septum in the axis.     

Observations of valve-like structures and evidence for rectification of flow within the gill lamellae of the crab Carcinus maenas (Crustacea,Decapoda)

Summary The isolated gills of Carcinus maenas, perfused at pressure drops of 1–10 cm of water, exhibited flow rectification, the resistance to perfusion via the afferent vessel being many times lower than that for efferent perfusion. The asymmetry was greater at the lower end of this pressure range.The overall afferent branchial resistance for Carcinus of weight 65 g, and with no ventilatory component in the transmural pressure difference, was estimated to be 0.05 cm of water. l^–1 · sec. The corresponding overall reverse (efferent) branchial resistance was 0.36 cm of water · l^–1 · sec.LM, TEM and SEM examination of the gills indicated that haemolymph leaves each gill lamella via several discrete parallel efferent channels which drain different regions of the lamella, and that each efferent channel is nearly closed, at its junction with the efferent branchial vessel, by a diaphragm of loosely interwoven and very elongated cells. It is concluded that these cells may constitute efferent valves and that narrow apertures between them may contribute a major component to the branchial resistance and be primarily responsible for the rectification of flow. Relatively wide apertures lead directly from the afferent vessel into the lamellae and are not asociated with valves of any kind.The valves may be important in enabling changes in transmural pressure associated with ventilatory reversals to pump haemolymph unidirectionally through the lamellae. Similarly valves may allow the oscillating venous pressures associated with locomotor activity to improve gill perfusion during exercise.The elongated tails of the cells of the efferent valve contain numerous microtubules. The wider cell bodies contain the nucleus and many mitochondria. Unusual organelles composed of many short (about 0.25 m long) microtubules and often lying close to the nuclear membrane may be microtubule organising centres. It is speculated that, in addition to their simple mechanical function, the valve cells may play a more dynamic role in regulating flow of haemolymph through different lamellar routes, or that they may monitor composition, pressure or flow of the efferent lamellar circulation.     

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.