Neuroendocrine Control Mechanisms in Shell Formation

The brain of Helisoma duryi contains several neurodendocrinecentres. Factors) present in the cerebral ganglia are thoughtto be involved in normal shell growth while neurosecretory substancespresent in the visceral ganglion are involved in the repairof damaged shell. In Lymnaea stagnalis a growth hormone is producedby the cerebral ganglion which stimulates periostracum formationand the calcification of the inner shell layer. The second effectis thought to occur through the action of a mantle edge calciumbinding protein. In Helisoma, mantle collar is able to produce the periostracumin vitro. The presence of brain from a fast growing donor increasesthe amount of periostracum produced by a mantle collar froma slow growing animal. This effect is further enhanced by theremoval of the lateral lobes. The periostracum produced by fastgrowing animals has a higher glycine content than that producedby slow growing snails. The presence of dorsal epithelial tissueenhances the incorporation of calcium into periostracum formedin vitro. These findings suggest that a single factor is present in thebrain of fast growing Helisoma which modulates shell formationrates in vivo and periostracum formation in vitro.     

REMOTE BIOMINERALIZATION IN DIVARICATE RIBS OF STRIGILLA AND SOLECURTUS (TELLINOIDEA: BIVALVIA)

Deposits composed of aragonite prisms, which were formed afterthe outer shell layer, have been found at the posterior steepslopes of divaricate ribs in two species of Strigilla and anothertwo of Solecurtus. These prisms have their axes oriented perpendicularto the outer shell surface and differ in morphology from fibresof the surface-parallel composite prisms forming the outer shell.They display crystalline features indicating that, unlike crystalsforming the outer shell surface, their growth front was free,unconstrained by the mantle or periostracum. These particulardeposits are called free-growing prisms (FGPs). In these generathe periostracum is clearly not the substrate for biomineralizationand, upon formation, does not adhere to the steep slope of ribs,but detaches at the rib peak and reattaches towards the posterior,just beyond the foot of the posterior scarps of ribs. In thisway, a sinus or open space developed between the internal surfaceof the periostracum and the outer shell surface along each steeprib slope. These spaces could remain filled with extrapallialfluid after the mantle advances beyond that point during shellsecretion. FGPs grow within this microenvironment, out of contactwith the mantle. Other species with divaricate ribs do not developFGPs simply because the periostracum adheres tightly to both ribslopes (which are never so steep as in Solecurtus and Strigilla).FGPs constitute one of the rare cases of remote biomineralizationin which aragonite is produced and direct contact with the mantlenever takes place. (Received 22 November 1999; accepted 20 February 2000)     

THE AMINO ACID COMPOSITION OF FRESHWATER MOLLUSC SHELLS IN RELATION TO PHYLOGENY AND ENVIRONMENT

Quantitative thin layer chromatography of structural amino acidswas standardized for the study of shell proteins of 13 speciesof freshwater mollusc. There was slight intraspecific variationbetween individuals, particularly for tyrosine concentrations.No external periostracum was chemically identifiable in theshell of Lymnaea peregra (Müller) and this species showedsome amino acid heterogeneity in different parts of the shell.Despite these intraspecific variations, analysis of variancesuggested interspecific variation in concentrations for almostall the amino acids tudied. Cluster and principal componentsanalysis indicated that the amino acid composition did reflectphylogenetic affinity but that environmental factors were probablymore-important. (Received 5 August 1982; revised 5 August 1982;     

MICROTUBULES IN THE SHELL OF THE INVASIVE BIVALVE CORBICULA FLUMINEA (MULLER, 1774) (BIVALVIA: HETERODONTA)

Using scanning electron and histological techniques on specimensof the bivalve Corbicula fluminea a new relationship betweenmantle, shell and periostracum has been observed, apparentlyfor the first time. Here we demonstrate that several extensionsof the mantle epithelium pierce the shell to join the innerlayer of the two-layered periostracum. The mantle extensionsare confirmed as unicellular processes. We suggest that theycould serve the animal in the mobilization of calcium from theshell for buffering the extrapalhal fluid under anaerobic conditions,when organic acids accumulate or when an extra contributionof Ca²⁺ is required (Received 5 January 1994; accepted 30 March 1994)     

Differentiation and Growth of the Brachiopod Mantle

Cell differentiation in the mantle edge of Notosaria, Thecidelhnaand Glottidia, representing respectively, the impunctate andpunctate calcareous articulate and chitinophosphatic inarticulatebrachiopods, is described. Comparison of electron micrographssuggests that outer epithelium which secretes periostracum andmineral shell, is separated from inner epithelium by a bandof "lobate" cells, of variable width, exuding an impersistentmucopolysaccharide film or pellicle. The lobate cells alwaysoccupy the same relative position on the inner surface of theouter mantle lobe; but the outer epithelium is commonly connectedwith the inner surface of the periostracum by papillae and protoplasmicstrands which persist during mineral deposition and ensure thatboth shell and attached mantle remain in situ relative to theoutwardly expanding inner surface of the outer mantle lobe.In the prototypic brachiopod, the lobate cells are likely atfirst to have occupied the hinge of the mantel fold but laterto have been displaced into their present position by the rigidoutward growing edge of the mineral shell.     

Boring of Shell by Caobangia in Freshwater Snails of Southeast Asia

Giard (1893) described Caobangia billeti, a freshwater sabellidpolychaete, based on material collected in what is now northernNorth Vietnam. Up to the present time it had not been re-collectedand Giard's observations that it is hermaphroditic, that itseggs undergo internal development, and that, rather than havinga posterior terminal anal opening, the anus opens far forward,have been questioned from time to time. Recently collected material, as well as older material in thecollections of the Division of Molluscs, Museum of Natural History,Smithsonian Institution, has allowed the confirmation ofGiard'sfindings and the observation of the burrows of Caobangia in22 species of molluscs from fast-moving streams in Ceylon, northernIndia, Assam, Burma, Thailand, Laos, Java, Sabah, and variousof the Philippine Islands. Burrowing in snail shells appears to take place subsequent tolarval metamorphosis as youngindividuals grow posteriorly froma capsule-like structure through the periostracum and into thecalcium carbonate of the snail shell. There appears to be nodifferentiated area of the worm which can be implicated as thesite of acidic secretions, and the setae of the worms seem notto be unusually worn, as would be the case if they were usedto wear away the shell material.     

THE ANATOMY OF CALLOCARDIA HUNGERFORDI (BIVALVIA: VENERIDAE) AND THE ORIGIN OF ITS SHELL CAMOUFLAGE

Callocardia hungerfordi (Veneridae: Pitarinae) lives in subtidalmuds (220 to 240m C.D.) and is covered by a dense mat of mudthat, effectively, camouflages the shell. The periostracum is two layered. The inner layer is thick andpleated, the outer thin and perforated. From the outer surfaceof the inner layer develop numerous, delicate (0.5 mm in diameter),calcified, periostracal needles. These penetrate the outer periostracum.Mucus produced from sub-epithelial glands in the inner surfaceof the mantle, slides over the cuticle-covered epithelium ofthe inner and outer surfaces of the inner fold and the innersurface of the middle mantle fold to coat the outer surfaceof the periostracum and its calcified needles. Increased productionat some times produces solidified strands of mucus which bindmud and detrital material into their fabric to create the shellcamouflage. Calcified periostracal needles have been identified in othervenerids, including some members of the Pitarinae, but how theyare secreted and how the covering they attract is producedand, thus, how the whole structure functions, has not been explained. (Received 7 December 1998; accepted 5 February 1999)     

THE FUNCTIONAL MORPHOLOGY OF THE ANTARCTIC BIVALVE THRACIA MERIDIONALIS SMITH, 1885 (ANOMALODESMATA: THRACIIDAE)

The functional morphology of the Thraciidae is poorly understood.Although some morphological aspects of several members havebeen described, only Trigonothracia jinxingae from Chinese watersis known in detail. Thracia meridionalis is the only representativeof the family in Antarctic waters, and is common in AdmiraltyBay, King George Island, where it inhabits muddy sediments.Thracia meridionalis shares many features with Tr. jinxingaethat are typical of most Anomalodesmata, i.e. a secondary ‘ligament’of thickened periostracum, extensively fused mantle margins,ctenidia of type E, a ctenidial-labial palp junction of categoryIII, a stomach of type IV and simultaneous hermaphroditism.Thracia meridionalis is, however, strikingly different fromTr. jinxingae in a number of ways, such as the presence of afourth pallial aperture, statocysts of type B₃, heterorhabdicctenidia, direct communication between the mantle chambers,a deep-burrowing habit (individuals lying on the left shellvalve), siphons that retract into mucus-lined burrows, a stomachwith extensive sorting areas, a rectum which passes over thekidneys and separate male and female gonadial apertures. Thereis, therefore, a greater range of morphological diversity withinthe Thraciidae than previously suspected. (Received 27 April 2004; accepted 30 November 2004)     

Phosphatic Shell Formation in Atremate Brachiopods

The atremate brachiopods are unique in that they possess shellsof calcium phosphate. In Lingula adamsi and Gloltidia pyramidata,the shell mineral is (CO₃ + F)-containing apatite and is crystallo-chemicallysimilar but not identical to the mineral francolite. The shellof Glottidia consists of a thin periostracum, a mineralizedthick primary layer, and alternating mineralized layers andless mineralized chitin layers. The basic unit of the crystalsis the spherulite. Proteinaceous and glycosaminoglycan (GAG)matrices are present in the primary and mineralized layers.The GAGS in the chitin layer are morphologically different fromthose of the other layers. The GAGS are intimately associatedwith the apatite crystals. Shell formation appears to be mediated by three different typesof cells in the outer epithelium. The cells primarily involvedin the mineral formation are characterized by many vacuoleswith electron-dense granular inclusions containing Ca, P, andS. The connective tissue at the anterior edge of the mantlealso contains fine granules with Ca, P, and S. Those granulesare considered to be a mineral reserve for shell formation.Some problems of the mechanisms of shell formation are discussed.     

淡水贝类贝壳多层构造形成研究

对几种淡水贝（包括蚌、螺）进行形态及组织学观察，并通过实验方法重现贝壳三种物质，即：角质、棱柱质、珍珠质的生成过程，结果表明：外套膜外表皮细胞是由相同类型细胞组成，这些相同细胞在不同的作用条件下形成贝壳多层构造。     

The Suitability of Some Algae for Mass Cultivation for Food, with Special Reference to Dunaliella bioculata

Twenty-five strains of freshwater and saltwater algae have beeninvestigated for their suitability for mass cultivation forfood. Under laboratory conditions a strain of Dunaliella bioculatahas been found to give yields comparable with those obtainedfrom Chlorella. It is suggested that only when the growing algalcells are exposed in very thin layers will the yield not belimited by light penetration. The storage polysaccharide ofD. bioculata contains a 1:4-glucosan resembling starch. Allthe essential amino-acids have been shown to be present withthe possible exception of methionine or valine and tryptophane.     

THE PALLIAL EYES OF CTENOIDES FLORIDANUS (BIVALVIA: LIMOIDEA)

The structure of the pallial eye in the Limidae has neverbeen elucidated properly, largely because they are difficultto see among the mass of surrounding mantle tentacles and becausethey are few, small, and lose their pigmentation when preserved.Possibly two eye types are present, simple cup-shaped receptorsin species of Lima, like those seen in the Arcoida, and morecomplex invaginated ones in Ctenoides. The pallial eyes (;18on both lobes) of Ctenoides floridanus are formed by invaginationof the middle mantle fold at the periostracal groove, so thatall its contained structures are derived from the outer andlight is perceived through the inner epithelia of this fold.The eye comprises a simple multicellular lens and a photoreceptiveepithelium beneath it of lightly pigmented cells and alternatingvacuolated, support cells. In some species of the Arcoidea, Limopsoidea and Pterioidea, pallialeyes occur on the outer mantle fold and thus beneath the periostracum(and shell). The pallial eyes of Ctenoides floridanus and otherpterioideans, e.g. species of the Pectinidae, occur on the middlefold and may thus have improved vision. In the Cardiodea, Tridacniidaeand Laternulidae (Anomalodesmata) pallial eyes occur on theinner folds. There is thus a loose phylogenetic trend, in which Ctenoidesis a critical link, of increasing eye sophistication correlatedwith the historical age of the clades possessing them. (Received 16 November 1999; accepted 20 January 2000)     

The Effect of Available Nitrate and Ammonium-Nitrogen on the Growth of Two Nitrogen-Fixing Blue-Green Algae

The effect of combined nitrogen supplied as nitrate or ammonium-nitrogenon the growth of two nitrogen-fixing blue-green algae, Nostocentophytum and Calothrix scopulorum, has been studied. Thesespecies have been isolated from marine environments. Both algaegrew as vigorously on elemental nitrogen as in the presenceof combined nitrogen. Growth was equal at all levels of nitrate-nitrogenemployed but high levels of ammonium-nitrogen proved inhibitoryor even toxic to the algae. Nostoc was slightly more susceptibleto high ammonium-nitrogen levels than was Calothrix. Increasein pH of the medium from 7.2 to 8.4 increased the toxic effectof ammonium-nitrogen although relative growth at the variouslevels of nitrate-nitrogen was not affected. The results suggestthat the different effects of ammonium-nitrogen on the growthof freshwater and marine blue-green algae may be due in partat least to the different pH levels of freshwater and marineenvironments in which the algae grow, rather than to any inherentdifference between the two groups.     

Osmotic and Ionic Regulation of North American Zebra Mussels (Dreissena polymorpha)

SYNOPSIS. Unlike other freshwater bivalves that survive formonths in deionized water, Dreissena polymorpha requires minimalconcentrations of Na, K, Mg, and Cl in the bathing medium forlong-term survival. Although ion transport rates are higherin D. polymorpha compared to other freshwater bivalves, theytend to have lower blood solute concentrations. D. polymorphahas an unusually "leaky" epithelium with a high paracellularpermeability to solutes. Thus, even with high transport rates,it may not be possible for zebra mussels to retain higher bloodsolutes because of the extensive passive loss of ions. Undera hyperosmotic stress, D. polymorpha will rapidly osmoconform(about 12 hr) due primarily to the diffusion of solutes andpartially to the osmotic loss of water. D. polymorpha is notcapable of surviving an imbalance of Na/K in the external medium.In the absence of K the cells will tend to lose volume to achieveisosmotic balance with the blood, but the animals usually diewithin a few days. If D. polymorpha is exposed to excess K inthe environment (1 mM), they will accumulate K in the blood.If the K enters the cells, cellular volume would expand dueto increase in osmolyte concentration, yet, if K remains inthe blood, there will be an electrochemical imbalance. In eithercase, the animal cannot survive much longer than a day. WhenNa and K are present in the medium in a balanced combinationapproximated by artificial seawater (ASW), D. polymorpha willsurvive an acute transfer to 100 mosm ASW indefinitely (months).Our preliminary studies have shown that D. polymorpha will toleratestep-wise acclimation to solutions >250 mosm provided thechanges in salinity do not exceed 50–100 mosm. Freshwaterbivalves, unlike the marine bivalves, have limited free aminoacids in their body fluids and must rely on inorganic ions forosmotic regulation. The free amino acids serve as an importantosmolyte buffer for volume regulation when an animal experiencesan environment of changing salinity. The inability of Dreissena,and perhaps other freshwater bivalves, to tolerate hyperosmoticallyinduced dehydration may be due, in part, to the inability toaccumulate or retain sufficient intracellular K to facilitateregulatory volume adjustments.     

Origin and evolution of a myxozoan worm

Buddenbrockia plumatellae is an active, muscular, worm-shapedparasite of freshwater bryozoans. This rare and enigmatic animalhas been assigned to the Myxozoa on the basis of 18S ribosomalDNA sequences and the presence of malacosporean spores. Herewe report cloning of four homologous protein-coding genes fromBuddenbrockia worms, the putatively conspecific sac-shaped parasiteoriginally described as Tetracapsula bryozoides and the relatedsac-shaped parasite Tetracapsuloides bryosalmonae, the causativeagent of proliferative kidney disease in salmonid fish. Analysesare consistent with the hypothesis that Buddenbrockia is indeeda malacosporean myxozoan, but do not provide support for conspecificitywith either T. bryozoides or T. bryosalmonae. Implications forthe evolution of worm-like body plans in the Myxozoa are discussed.     

Function of the flexible periostracal hairs in Trichotropis cancellata (Mollusca, Gastropoda)

Abstract. The marine snail Trichotropis cancellata possesses hairy projections of periostracum (outer shell layer) whose function is unknown. Although rigid shell projections in molluscs have been studied extensively, the selective advantage of flexible extensions of periostracum is less clear. None of the functions proposed previously for periostracum (e.g., protection from erosion and boring) are promoted when it is drawn into hair-like projections. We investigated hypothetical functions that may be served by flexible periostracal hairs, including predator deterrence, alteration of flow vectors to promote feeding or affect turbulence dynamics during freefall, and providing a differential substratum for epibionts. Our laboratory results indicate that crabs, Cancer oregonensis , and sea stars, Pycnopodia helianthoides , consumed snails with the periostracum removed more often than snails with an intact hairy periostracum. However, in both predatory species, some individuals showed no significant preference, and another crab species ( Cancer productus ) did not strongly discriminate based on the shell periostracum. Field studies showed no difference in the rate of predation on hairy- versus smooth-shelled snails. The hairs did not alter flow around the shells consistently in laboratory flume experiments. Additionally, hairy- and smooth-shelled kleptoparasitic snails grew at rates that were statistically indistinguishable, while hairy, suspension-feeding snails grew more slowly. The hairs did not impact the orientation of a snail after a falling event or the time to righting after a fall. The presence of the hairs did deter settlement by barnacles. We conclude that the hairy periostracum acts as a slight deterrent to crab and sea star predators and as a stronger deterrent to the settlement of large calcareous epibionts, such as barnacles, that would increase the weight the snail must bear and potentially increase drag.     

Culturing freshwater, planktonic rotifers on Rhodomonas minuta var. nannoplanctica Skuja and Stichococcus bacillaris Nageli

Eleven freshwater, planktonic rotifer species (Brachionus angularis,Brachionus cafyciflorus, Keratella quadrata, Nolholca squamula,Ascomorpha ecattdis, Synchaeta kitina, Synchaela tremula, Potyarthradolichoptera, Polyanhra vulgarii, Pompholyx sulcata, Filiniabrachiata) were cultured successfully on either Rhodomonas minutavar. nannoplanctica Skuja or Stichococcus bacillaris Nägeli,in a totally synthetic medium. Three of these species had notpreviously been kept in laboratory culture. The method is proposedas a simple technique for the routine batch culture of a varietyof planktonic rotifer species. Some important observations onhandling techniques, incubation temperature, culture vesselsand media are discussed.     

A hairy business—Periostracal hair formation in two species of helicoid snails (Gastropoda,Stylommatophora, Helicoidea)

In molluscs, the calcareous shell is covered externally by a thin organic layer, the periostracum. The periostracum of some pulmonate species is of special taxonomic interest because it bears distinct microscale architectures. Where and how these structures are formed is as yet unknown. Using histological sections through their shells, gelatin cuts, and live observations I studied the pattern by which the periostracal hair‐like projections in two helicoid land snail species are secreted and evenly arranged on the shell. The results indicate a complex mechanism: a hair is formed in the periostracal groove independently of the periostracum, after which it is attached to the edge of the shell, drawn out of the tissue, and finally swivelled to the upper side of the periostracum. Upon further growth of the periostracum, the hairs are finally fixed upright on the shell. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Cortisol and Pacific Salmon: A New Look at the Role of Stress Hormones in Olfaction and Home-stream Migration

Pacific salmon (genus Oncorhynchus) exhibit an interesting anduncommon life-history pattern that combines semelparity, anadromy,and navigation (homing). During smoltification, young salmonimprint on the chemical composition of their natal stream water(the home-stream olfactory bouquet or "HSOB"); they then migrateto the ocean where they spend a few years feeding prior to migratingback to their natal freshwater stream to spawn. Upstream migrationis guided by the amazing ability to discriminate between thechemical compositions of different stream waters and thus identifyand travel to their home-stream. Pacific salmon demonstratemarked somatic and neural degeneration changes during home-streammigration and at the spawning grounds. The appearance of thesepathologies is correlated with a marked elevation in plasmacortisol levels. While the mechanisms of salmonid homing arenot completely understood, it is known that adult salmon continuouslyutilize two of their primary sensory systems, olfaction andvision, during homing. Olfaction is the primary sensory systeminvolved in freshwater homing and "HSOB" recognition, and willbe emphasized here. Previously, we proposed that the increasein plasma cortisol during Pacific salmon home-stream migrationis adaptive because it enhances the salmon's ability to recallthe imprinted memory of the "HSOB" (Carruth, 1998; Carruth etal., 2000b). Elevated plasma concentrations of cortisol couldprime the hippocampus or other olfactory regions of the brainto recall this memory and, therefore, aid in directing the fishto their natal stream. Thus, specific responses of salmon tostressors could enhance reproductive success.