Comparative Functional Morphology of the Bivalve Excretory System

Combining injection techniques with ultrastructural observations,and relating these findings to the more traditional physiologicaland morphological studies have shed new light onthe excretorymechanisms underlying the processes of ultrafiltration, secretionand reabsorption in some bivalve molluscs. These basic processesare further elucidated by comparing normal excretory tissueswith those in bivalves that have been subjected to stress bypollutants in either the natural environment or under laboratoryexperimentation. The process of ultrafiltration is size andcharge dependent and occurs at the filtration barrier at thebase of the podocytesin the pericardial gland. Primary urinemay be modified by secretion (primarily from the kidney cellsbut also from the podocytes), reabsorbtion in the kidney, andby the addition of hemocytes passing from blood spaces throughthe epithelium into the lumen of the kidney. Numerous concrements(granules, concretions and membranes) that result from lysosomalactivities inthe podocytes, kidney cells and hemocytes alongwith the fluid are excreted into the mantle cavity     

The effects of temperature on the distribution and establishment of Echinoparyphium recurvatum metacercariae in Lymnaea peregra

The establishment and distribution of Echinoparyphium recurvatum metacercariae in the second intermediate host, Lymnaea peregra, were investigated at a temperature range of 5-29 degrees C. Preliminary studies on the survival and infectivity of E. recurvatum cercariae showed that both parameters were temperature dependent. No cercarial transmission occurred at 5 or 10 degrees C. Nevertheless, the transmission efficiency (1/H0) indicated that transmission was temperature independent in the temperature range 17-25 degrees C and was much lower than in previous studies using this host-parasite system. These differences were attributed to low cercarial densities used in this study. The effect of temperature on encystment site choice (mantle cavity, kidney, pericardium) by metacercariae showed that the mantle cavity was the prime site of encystment, followed by the pericardium and the kidney. Temperatures at the lower and upper ranges (14 and 29 degrees C), however, caused a significant reduction in encystment in the mantle cavity but not in the pericardium or kidney. The importance of cercarial densities, the physiological mechanisms influencing metacercarial distribution and their implications for parasite transmission to the definitive host are discussed.     

Fine structure of the kidney and characterization of secretory products inDentalium rectius (Mollusca,Scaphopoda)

Summary The ultrastructure of the scaphopod kidney and secretory product composition is described, for the first time, inDentalium rectius. The kidney epithelium consists of two primarily secretory cell types. The first exhibits extensive vacuolation, and scattered granules are formed within the vacuolar space by a process of surface accretion; the incorporation of glycogen particles in this process is associated with very fine, electronopaque threads which radiate from the granule. The second cell type possesses granules enclosed individually within secretory vesicles, and intermediate stages in their growth are characterized by needle-like crystals on the granule surface. The secretory vesicles in some cases coalesce to form a large central vacuole filled with granules. This cell type possesses an apical membrane with sparse microvilli, which may indicate a secondary reabsorptive capacity. Granules in both cell types show a concentric ring ultrastructure, and are composed primarily of calcium phosphate with a small amount of zinc; there is also an organic component of protein, mucopolysaccharide and a large amount of glycogen. Ultrastructural and histochemical observations indicate a lysosomal origin for the granules, although granules of the second cell type develop intracellularly to a greater extent than those of the first. All granules are extruded into the kidney lumen by a process of merocrine secretion prior to release into the mantle cavity via an externally ciliated, muscular excretory pore.     

The influence of mantle cavity fluid on the aerial oxygen consumption of some intertidal gastropods

The intertidal gastropods Gibbula cineraria (L.), Nucella lapillus (L.), and Littorina littorea (L.) have been investigated. Animals that had attached to a plastic surface under sea water were exposed to air and rapidly frozen. Fluid was found to be retained within the mantle cavity in contact with the gill. For each species the weight of the mantle cavity fluid retained in air was related to the whole weight of the animal. There was no significant difference between the weight of fluid held by animals in the laboratory and that held by animals on the shore 2 h after aerial exposure. The oxygen consumption of inactive individuals in air was measured by two techniques which did not produce significantly different results. The aerial oxygen consumption of all three species was lowered by the loss of mantle cavity fluid. This also reduced the temperature coefficient. A correlation is suggested between a large weight of mantle cavity fluid, the presence of a gill in the mantle cavity and a large reduction in oxygen consumption upon the loss of fluid.     

Development of asymmetry in the caenogastropods Amphissa columbiana and Euspira lewisii

Abstract. The asymmetry displayed by the body plan of gastropods has been directly or indirectly attributed to an evolutionary process called torsion. Torsion is defined as a rotation of 180° between the cephalopodium (head and foot) and visceropallium (visceral organs, mantle, mantle cavity, and shell). During development, the displacement of anatomical components occurs during a process called "ontogenetic torsion." Although ontogenetic torsion is central to theories of gastropod evolution, surprisingly few studies have documented actual tissue movements during the development of asymmetry in gastropods. We investigated the development of the mantle cavity and pleurovisceral nerve connective (visceral nerve loop) in the caenogastropods Amphissa columbiana and Euspira lewisii , because displacements of both of these structures are interpreted as major consequences of torsion. Scanning electron micrographs, histological sections, and immunofluorescence images showed that the developing vis-ceropallium twists by 90° relative to the cephalopodium, the mantle cavity initially forms on the right side, and displacements of the visceral nerve loop become evident on the left side before the right side. A developmental stage in which the mantle cavity is confined to the right side has also been reported in members of the Vetigastropoda and Heterobranchia. We suggest that further comparative studies should test the hypothesis that early development throughout the Gastropoda converges on an embryonic organization in which the mantle cavity and anus are located laterally, despite clade-specific differences in developmental patterns both before and after this stage.     

Structure and function in the excretory systems of some terrestrial prosobranch snails (Cyclophoridae)

The excretory systems of terrestrial prosobranch snails of the family Cyclophoridae, collected in Jamaica, Costa Rica and South Africa, have been examined physiologically and as regards their gross and fine structure. The process of urine formation commences in the heart, where fluid is filtered across the wall of the ventricle. Filtration through the auricular wall is believed to be negligible. The kidney, which contains three types of cell, modifies the composition of the filtrate. One type of resorptive cell, characterized by basal infoldings associated with mitochondria, takes up salts. Another type, with basal subcellular spaces, may be responsible for taking up water. The third type of cell is secretory, producing concretions of uric acid and phospholipid which are liberated into the kidney lumen when the cell degenerates.
The rate and mechanism of urine production have been investigated using injections of inulin. The filtration rate at 25°C is 0.5 μl/g/min, and in 100% R.H. the average rate of urine production is 0–39 μl/g/min.
An accessory excretory organ has been developed from the hypobranchial gland of aquatic forms. It is composed of groups of subepithelial tubular glands opening into the mantle cavity by one or a series of pores, and secreting purines, phospholipids and mucus. There is evidence that this organ becomes progressively more complex in forms occupying drier habitats.
The systems of excretion and osmoregulation in the Cyclophoridae are considered to be very similar to those in their aquatic relatives, the Viviparidae and Ampullariidae. Certainly the cyclophorids are not as well adapted to a terrestrial life as are the Pulmonata, and in many respects they may be considered "aquatic" snails living on land.     

Neuroeffector connections of multimodal neurons in the African snail (Achatina fulica)

Using a new method of animal preparation, the efferent connections of giant paired neurons on the dorsal surface of visceral and right parietal ganglia of snail, Achatina fulica, were examined. It was found that spikes in giant neurons d-VLN and d-RPLN evoke postjunctional potentials in different points of the snail body and viscerae (in the heart, in pericardium, in lung cavity and kidney walls, in mantle and body wall muscles, in tentacle retractors and in cephalic artery). The preliminary analysis of synaptic latency and facilitation suggests a direct connections between giant neurons and investigated efferents.     

Morphology and biology of Polysaccus japonicus (Crustacea, Rhizocephala, Akentrogonida, Polysaccidae, fam. n.), a parasite of the ghost-shrimp Callianassa japonica

The morphology and biology of Polysaccus japonicus Høeg & Lützen, 1993 is described. Nearly all infected hosts are adults, the females of which become castrated. Each bears 4–50 (female) externae on the underside of the abdomen. The externae are interconnected via a root system within the host's perineural blood sinus. The externae pass through three moults. Cypris larvae inject spermatogonia into the immature externa's mantle cavity, which is where spermiogenesis occurs later. Multiplication of the male cells is probably supported by secretion from paired mantle cavity glands, presumably homologous to the receptacles of kentrogonid rhizocephalans. Late during the 2nd intermoult the visceral mass ruptures to release the ova into the mantle cavity where they become fertilized. The third moult results in opening of the mantle aperture which initiates a period of vigorous peristalsis of the muscular mantle. Following emission of the cypris larvae the externae perish leaving distinct scars. The root system always contains buds of various sizes and stage of development. When the scarred host moults, the most advanced buds emerge to give rise to a new generation of externae. This is thought to be repeated for the rest of the host's life.
The antennule of the cyprid has a relatively very long and slender 3rd segment, a unique construction among Akentrogonida. Because of this and the presence of special mantle cavity glands, a new family, Polysaccidae, is erected, which in some characters is intermediate between the Kentrogonida and the Akentrogonida.     

Larval development and metamorphosis in the marine pulmonate Amphibola crenata (Mollusca: Pulmonata)

The development of the free-swimming veliger of Amphibola is followed from hatching to settlement, and the larval structures compared with those of post-metamorphic juveniles and adult snails. Observations of living specimens and light-microscope sections were combined with scanning electron microscopy to build up a composite picture of veliger structure.
Four stages in the development of veligers are recognized, each being characterized by the appearance of organ systems such as the mantle cavity, larval heart, adult heart and kidney, and larval pallial gland. At or after metamorphosis, the larval systems (heart, kidney and pallial gland) disappear, and the developing adult organs move to the positions characteristic of adult snails.
Organogenesis in Amphibola veligers is compared with that of prosobranch and opisthobranch larvae, and with that of pulmonate larvae with direct development. The closest similarity is seen to be with opisthobranch veligers.     

Deposit feeding in Abra tenuis (Bivalvia: Tellinacea)

Deposit feeding in Abra tenuis is described in terms of the size of particles utilized.
Material is collected by the inhalant siphon performing a circular motion sucking in sediment from beneath and on the surface.
The size distribution of silica admitted into the mantle cavity is described and shown to be controlled by physical parameters. The density of a particle does not affect its uptake by the inhalant siphon. The size distribution of the sediment affects the size distribution of particles admitted to the mantle cavity.
No selection of material for ingestion in terms of size occurs after it has been taken into the mantle cavity. Thus the range of material ingested is ultimately controlled by a physical parameter, the inhalant opening.     

The excretory function of the posteriormost part of the echinoid and holothuroid gut (Echinodermata)

In vivo and in vitro experiments demonstrate that the intestinal caecum of the echinoid Echinocardium cordatum and the rectum of the holothuroid Holothuria tubulosa actively excrete chlorophenol red (CPR) from the individual body cavity towards the gut lumen, a function already demonstrated for the rectal caeca of asteroids. Physiological and ultrastructural investigations of the enterocytes of these organs show that they are functionally and structurally analogous to the cells of the proximal tubules of the vertebrate kidney. The results allow to suggest that the capability to eliminate unwanted substances through the posteriormost part of the gut is a generalized feature within the Echinodermata.     

Morphometric parameters of nutria kidney structures in postnatal ontogeny

Data on the morphometric parameters of the renal corpuscle, renal tubules, and collecting ducts of male and female nutrias in postnatal ontogenesis were obtained. It was found that the area of the renal corpuscle, glomerulus, the cavity and lumen of the capsule, and the proximal tubule diameter in the right and left kidney of female and male nutrias in the first year of life increase. The distal tubule diameter also increases; however, the dynamics of its changes becomes sinuous after 4.5 months. The collecting duct diameter varies depending on gender, age, and renal topography. The nuclear-cytoplasmic ratio in the cells of proximal and distal tubules and collecting ducts changes in a sinuous manner and depends on the gender and age of nutrias. The minimum mean value of the nuclear-cytoplasmic ratio was found in the proximal tubule cells in the left kidney of 12-month-old female nutrias (0.162 ± 0.002), and the maximum value was found in the distal tubule cells in the left kidney of newborn male nutrias (0.435 ± 0.007).     

Fine structure of the prothoracic mycangium, a chamber for the culture of symbiotic fungi, in the southern pine beetle, Dendroctonus frontalis

The ultrastructure of the prothoracic mycangium of female Dendroctonus frontalis is examined. The mycangium consists of a cuticular invagination within which symbiotic fungi are cultured by the pine beetle and transported to new host trees. Secretions from two types of gland cells pass into the mycangial lumen. The plasma membrane of type-1 cells is invaginated to form an enclosed extracellular cavity. The secretory product passes into the cavity, then through fine cylindrical channels into an end apparatus and finally via an efferent cuticular ductule to the lumen of the mycangium. Secretion of the type-2 cells is released into a cavity just beneath the mycangial cuticle. The cuticle over this cavity is quite thin (1-2mu), consisting mostly of inner epicuticle riddled with irregular canals through which the secretion reaches the lumen. Beneath the patches of porous cuticle are ribs (up to 1Omu in thickness) which flank the cavities and presumably provide structural support for the porous secretory zones.     

A new species of Tetrastemma (Nemertea: Enopla: Monostiliferoidea) from Ria de Foz, north-western Spain, found living in the mantle cavity of the bivalve mollusc Scrobicularia plana

A new species of monostiliferoidean nemertean, Tetrastemmafozensis , from the mantle cavity of the bivalve mollusc Scrobicularia plana , in the Ría de Foz, north-western Spain, is described and illustrated. Other nemerteans from the mantle cavity of Scrobicularia collected in Poole Harbour, southern England, are provisionally identified as conspecific with the Ria de Foz material, but show certain anatomical differences whose taxonomic significance cannot at present be assessed.     

黄鳝的泌尿系统及其功能

黄鳝腹腔左侧有盲端的中空管状结构,不是退化性腺,而是十分特化的长管囊膀胱。膀胱内壁具大量发达的绒毛,绒毛表面是移行上皮。在绒毛内部或基部有丰富的血管,因此该管囊膀胱不仅可贮存尿液,而且可能对水分等有重吸收作用。在中肾管与膀胱相接处,膀胱腔背侧出现一条明显的纵行皱襞,即在其横切面上观为巨绒毛。巨绒毛形成的原因,主要是中肾管移入膀胱壁所致。中肾管在巨绒毛内移行一程后才开口于膀胱。因此,黄鳝的生殖腺不是一对,而是一个,位于腹腔右侧。黄鳝肾脏细长,呈“丫”,在腹腔背侧,紧贴脊椎。前端为头肾,无肾单位,仅是造血器官。中肾有类似于哺乳类的肾小体,但数量较少,主要分布在肾脏周缘。肾小管包括颈段、初级近曲小管、次级近曲小管、初级远曲小管和次级远曲小管。两中肾管位于两肾叶腹内侧,其上皮间或是假复层柱状上皮,间或是移行上皮。前者含有许多杯状细胞,并可见到顶浆分泌的现象。中肾的肾小管间组织是大量的红细胞样组织,头肾似具有贮存或释放刚成熟的红细胞的组织结构,因此黄鳝肾脏可能是体内主要的造血器官。     

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.     

Water balance and the mantle cavity fluid of Nucella lapillus(L.) (Mollusca: Prosobranchia)

When uncovered by the tide Nucella lapillus (L.) retains fluid in the mantle cavity which accounts for ≈ 39% of the total water content of the attached animal. This extra-corporeal fluid reservoir retained in the mantle cavity has been investigated in relation to desiccation. Volume changes of mantle cavity fluid (m.c.f.) and concentration changes in Cl^?, Na⁺ and total osmotic pressure of m.c.f. and blood, suggest that the m.c.f. reduces the full effects of desiccation on the animal by restricting those effects to the external compartment. The blood of Nucella living in normal sea water was found to be hypo-osmotic and an hypothesis is presented which links this to the rôle of the m.c.f. during desiccation.     

Modern insights on gastropod development: Reevaluation of the evolution of a novel body plan

More than a century of speculation about the evolutionary originof the contorted gastropod body plan has been inspired by adultanatomy and by long-standing developmental observations. Theresult has been a concept of gastropod torsion that I call the"rotation hypothesis." Under the rotation hypothesis, gastropodsoriginated when all components of the visceropallium (shell,mantle, mantle cavity with contained structures, and viscera)rotated by 180° relative to the head and foot. This evolutionaryrotation is echoed during early development of patellogastropodsand vetigastropods and occurs to some extent during developmentof more derived clades. However, comparative developmental dataon ontogenetic torsion are minimal and I argue that the rotationhypothesis is a tautological argument. More recent studies onrepresentatives from 3 major clades of gastropods suggest thatthe highly conserved aspect of gastropod development is notsynchronous rotation of all components of the visceropalliumrelative to the head and foot but rather a state of anatomicalorganization in which the developing mantle cavity is on theright but the shell coil is posterior (endogastric orientation).This conserved state of developmental anatomy has inspired analternative hypothesis for the evolutionary origin of the gastropodbody plan, the "asymmetry hypothesis." Under the asymmetry hypothesis,the gastropod mantle cavity originated from 1 side only of abilateral set of mantle cavities. The asymmetry hypothesis doesnot require a saltation event to explain the origin of gastropods,nor does it require that the ancient molluscan precursor ofgastropods carried the shell coil over the head (exogastricorientation).     

Embryonic development and organogenesis in the snail Marisa cornuarietis (Mesogastropoda: Ampullariidae). V. Development of the nervous system.

The nervous system is ectodermal in origin. All nerve ganglia arise separately by proliferation and later delamination from the ectoderm, not by invagination. They become secondarily connected to one another by commissures and connectives developing as extensions from the peripheral layer of ganglionic nerve cells. Rudiments of the cerebral, pedal, pleural and intestinal (parietal) ganglia arise almost simultaneously at a relatively early stage (Stage V). The cerebral ganglia develop from the ectoderm of the head plates. Rudiments of the pedal and pleural ganglia are separate at their inception. They later fuse (Stage VI) to form a pleuro-pedal ganglionic mass on each side. The 2 intestinal ganglia are symmetrical at the beginning, but they soon lose their symmetry as a result of torsion. The right ganglion crosses to the left over the gut and persists as the supraintestinal ganglion. The left or subintestinal ganglion shifts to the right and forward, and fuses with the right pleural ganglion (Stage VIII), thus obscuring the chiastoneury. The paired buccal and single visceral (abdominal) ganglia start differentiating in Stage VII. The former develop from the ectodermal wall of the stomodaeum, while the visceral ganglion delaminates from the right wall of the visceral sac, then shifts to the left during torsion. The statocysts develop early (Stage V) from 2 ectodermal invaginations on either side of the rudimentary foot. They later separate from the overlying ectoderm and statoconi appear in their lumina. Contrary to earlier reports on related ampullariids, the osphradium proved to be ontogenetically older than the mantle and mantle cavity. It starts differentiating as a thickened ectodermal plate in the right wall of the visceral sac (Stage V). During torsion, it becomes engulfed in the mantle cavity and shifts to the left side, then is carried forward as the mantlegrow. The eyes develop late (Stage IX) as ectodermal invaginations which rapidly separate from the ectoderm to form closed vesicles. Their cells start differentiating before hatching to form the retina, in which pigment is deposited, and the inner cornea. The lens is secreted in the lumen of the eye and grows by addition of concentric layers of secretion.