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)     

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.     

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 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)     

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.     

PREDATION ON THE BIVALVE CHOROMYTILUS MERIDIONALIS (KR.) BY THE GASTROPOD NATICA (TECTONATICA) TECTA ANTON

The effects of a population of the boring gastropod Natica tectaon the bivalve Choromytilus meridionalis were investigated atBailey's Cottage, False Bay, South Africa. In July 1979 theN. tecta density on the mussel bed averaged 69 m^–2 andthe population consisted mainly of reproductively mature individualsbetween 20–33 mm shell width. Laboratory experiments on N. tecta showed that prey size selectionis an increasing function of predator size. The prey size rangetaken by large N. tecta is also greater than that taken by smallindividuals. The position of the borehole on the mussel shellis a function of the way in which the shell is held by the footduring the boring process. Consumption rates measured in thelaboratory showed an increase from approximately 1 kJ per weekin 18 mm N. tecta to 4.5 kJ per week in 28 mm individuals. Populationconsumption in the field was calculated as 663 kJ m^–2month^–1. It was estimated that at this rate the standingcrop of mussels in the pool would be eliminated within 10 months.Field measurements showed significant depletion after 6 months. New spat settlement of mussels occur every 4–6 years.The growth curve shows that after one year the population meansize exceeds 30 mm shell length, which is beyond the prey selectionsize range of small N. tecta. It was concluded that at the timeof a new mussel settlement a niche is provided for the simultaneoussettlement and growth of juvenile N. tecta in high densities.However, within one year the increase in prey size, togetherwith depletion due to over-exploitation, limits population growthand density in N. tecta. (Received 14 March 1980;     

Studies on the in vitro formation of periostracum inHelisoma duryi: the influence of brain

Summary Excised mantle tissue produces periostracum when placed in organ culture. Mantle collars taken from animals exhibiting a fast rate of shell deposition in vivo produce more periostracum than mantle collars from slow growing animals. The addition of a brain from a fast growing animal increased periostracum production by a mantle collar from a slow growing animal. This effect was enhanced by removing the cerebral ganglia lateral lobes. This suggests that a factor(s) is present in the brain of fast growing animals which enhances periostracum formation in vitro. The lateral lobes appear to inhibit this activity. Radiolabel incorporation studies suggest that the periostracum produced by fast growing animals has a higher glycine content than that produced by slow growing animals.Abbreviations Dopa 3,4-dihydroxyphenylalanine - FB brain derived from a fast growing animal - FMC mantle collar derived from a fast growing animal - LL lateral lobes - LL lateral lobes removed - MC mantle collar - SB brain derived from a slow growing animal - SGPF shell growth promoting factor - SMC mantle collar derived from a slow growing animal - TCA trichloroacetic acid - WM whole mantle     

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)     

GROWTH OF JUVENILE BITHYNIA TENTACULATA (PROSOBRANCHIA, BITHYNIIDAE) UNDER DIFFERENT FOOD REGIMES: A LONG-TERM LABORATORY STUDY

The growth of juvenile Bithynia tentaculata (Proso-branchia,Bithyniidae) was measured under controlled laboratory conditionsover 18 months. The animals were fed with different concentrationsof suspended food (Chlamydomonas reinhardii at 2, 000 cells.ml^–1and at 10, 000 cells.ml^–1), solid food (lettuce) and combinationsof both (lettuce with 2, 000 cells.ml^–1 and with 10, 000cells.ml^–1 Chlamydomonas). The growth of all animals wasmeasured weekly, and after 1 years final shell sizes, shelldry weights and ash-free dry weights, as well as soft body dryweights and ash-free dry weights were determined. Survival rateof the animals increased from 20% when fed with 2, 000 algalcells.ml^–1 to 75% with lettuce and 10, 000 cells.ml^–1as food. Final shell sizes and shell weights were not influencedby food, but soft body dry weights were significantly reducedwhen animals were fed on suspended food only. Reproduction (i.e.egg laying) was observed at the end of the second summer whenlettuce (with or without algal addition) was given as food.The ecological consequences of these results are discussed. (Received 7 April 1993; accepted 9 August 1994)     

REPRODUCTIVE CYCLE AND FECUNDITY OF THE DATE MUSSEL LITHOPHAGA LITHOPHAGA (BIVALVIA: MYTILIDAE)

The reproductive cycle and fecundity of the date mussel Lithophagalithophaga, L. a well-known, ediblespecies has been examined.Sexes are separate and the mean number of eggs per each gonadis 1.894 x 10⁶ ± 1.044 x 10⁶ S.D. Reproduction firstoccurs at an age of 2⁺years and at a shell length greater than0.9 cm. Gonad activity is annual and is observed at all ages.The sex ratio for individuals up to 7 cm is 3:1 in favour ofthe males, whereas it becomes 1:1 for individuals greater than7 cm. The release of gametes by males and females occurs almostsimultaneously and begins immediately after a decline in thehighest water temperature ( 27°C), an increase in salinity(>31) and a decrease in the dissolved oxygen (6.5 ppm). Smallpercentages of mature individuals appear during the first wintermonths thus lengthening the reproductive period. This phenomenonis attributed to the temperature difference in deeper waters,the delay in gamete release by young individuals, tide, waveaction and changes in salinity. The fecundity of Lithophagalithophaga is high because its life-cycle is adversely affectedby environmental factors such as waves and tides. Fecundityincreases with shell length, more so with total wet weight andmainly with the age of the animals. Summer seems to be the suitableseason to exploit the date mussel of shell lengths > 5 cm. (Received 6 September 1993; accepted 10 March 1994)     

A new model for periostracum and shell formation in Unionidae (Bivalvia, Mollusca)

The periostracum in Unionidae consists of two layers. The outer one is secreted within the periostracal groove, while the inner layer is secreted by the epithelium of the outer mantle fold. The periostracum reaches its maximum thickness at the shell edge, where it reflects onto the shell surface. Biomineralization begins within the inner periostracum as fibrous spheruliths, which grow towards the shell interior, coalesce and compete mutually, originating the aragonitic outer prismatic shell layer. Prisms are fibrous polycrystalline aggregates. Internal growth lines indicate that their growth front is limited by the mantle surface. Transition to nacre is gradual. The first nacreous tablets grow by epitaxy onto the distal ends of prism fibres. Later growth proceeds onto previously deposited tablets. Our model involves two alternative stages. During active shell secretion, the mantle edge extends to fill the extrapallial space and the periostracal conveyor belt switches on, with the consequential secretion of periostracum and shell. During periods of inactivity, only the outer periostracum is secreted; this forms folds at the exit of the periostracal groove, leaving high-rank growth lines. Layers of inner periostracum are added occasionally to the shell interior during prolonged periods of inactivity in which the mantle is retracted.     

INTRA-POPULATION VARIATION OF THE SHELL OF LITTORINA RUDIS (MATON) (MOLLUSCA: PROSOBRANCHIA)

Shell variation within a single population of Litlorina rudis(Maton), collected near the Biological Station at Trondheim,Norway, was studied. The variation of the shell features studied, which includedthe shape of the basal part of the outer lip, and the relativeheight of the spire, width of the shell and of the aperture,were found to be at least partially related to shell height.Furthermore, the shell shape in the population investigated,besides varying with age, also varies due to shell damage. Nosignificant differences were found between the shapes of theshells of males and females. The shell characteristics of different populations of L. rudisalso vary greatly, one of the causes being differences in theenvironmental conditions. When studying this variation betweendifferent populations, however, it must be emphasized that onlyundamaged shells of individuals of about the same age shouldbe compared, in order to obviate the effects of the variabilitydue to age differences and shell damage within each individualpopulation. ^*Contribution from Trondhjem Biological Station no. 199 (Received 18 February 1980;     

ASPECTS OF THE LIFE CYCLE, POPULATION DYNAMICS,GROWTH AND SECONDARY PRODUCTION OF THE EDIBLE SNAIL HELIX LUCORUM LINNAEUS, 1758 (GASTROPODA, PULMONATA) IN GREECE

This paper examines the biology and ecology of Helix lucorumL. which lives in mainland Greece, as well as its growth andsecondary production. A demographic study revealed that (a)3 cohorts exist at any time during the year (when adults ofall generations belong to the same cohort) (b) egg-laying andhatching occur during the months of July and August respectively,(c) the most rapid growth takes place during spring. Study ofH. lucorum genitalia in relation to age showed that the snailsaxe sexually mature 3 years after hatching, when the largestdiameter of their shell (D) is equal to or greater than 35 mm Von Bertallanffy's method suggests that Helix lucorum may liveup to 14 years or more in order to reach its possible maximumsize (48.80 mm) The study of relative growth of D in relation to P_s (peristomesurface) of Helix lucorum shows that D grows faster than P_swhen D12.50 mm; juveniles change their growth rate when theirD arrives at 22.05 mm, and growth becomes slower when adultsarrive at 36.27 mm Annual secondary production calculated by the size frequencymethod gave a mean annual density of 3.39 individuals per m²,a mean annual crop (biomass) of 4.04 g-m^–2 and an annualproduction (P) of 5.02 g · m^–2. The annual turnoverratio (P//b) is equal to 1.24 (Received 23 June 1987;     

ASPECTS OF THE BIOLOGY OF THE GIANT FORM OF STHENOTEUTHIS OUALANIENSIS CEPHALOPODA: OMMASTREPHIDAE) FROM THE ARABIAN SEA

An investigation was carried out on the recently discovered‘giant’ extra large (XL) form of the squid Sthenoteuthisoualaniensis from the Arabian Sea. The sample consisted of 2males, which have not been previously described, and 13 females.Diet composition, parasite loading, sucker ring dentition, biolumi-nescenceand sexual dimorphism were examined and compared to known parametersof the medium (M) form. Reproductive strategy, potential fecundity,egg size distribution in the ovary and oviducts were examinedin mature XL females. Evidence of multiple spawning in the giantform was also investigated. Overall body shape, bioluminescentstructures and coloration of the giant form were similar tothe M form, though the XL form had a smaller fin angle thanthe M form. The mature female XL form has a dorsal mantle lengthabout twice that of a mature female M form. Adult females ofthe XL form have a dorsal mantle length about twice that ofadult males of the same form. Differences between males andfemales were found in arm sucker ring dentition and parasiteload, suggesting a difference in diet. This could be linkedto size differences between the sexes. A strong correlationbetween ovary mass and mantle length was found (r² = 0.64).Poor correlation was found between mantle length and oviductmass (r² = 0.128) and potential fecundity (r² = 0.07). Potentialfecundity ranged between 2–5 million eggs and the holdingcapacity of the oviducts was approximately 300, 000 eggs. Thiscombined with the presence of spermatangia and the presenceof food in the stomach suggest that the XL form is a multiplespawner. S. oualaniensis appears to have a plastic phenotypeand has adapted to the Arabian Sea conditions by evolving thecapacity to grow to a giant size. (Received 6 November 1996; accepted 15 February 1997)     

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 MANTLE AND SHELL OF SOLEMYA PARKINSONI (PROTOBRANCHIA: BIVALVIA)

The shell of Solemya exhibits considerable flexibility which is further enhanced by the marked extension of the periostracum beyond the calcareous portions of the valves. This fcature, more than any other, has made possible the habit, unique among bivalves, of burrowing deep within the substrate without direct contact with the water above. The inner calcareous layer of tho valves is restricted to a small area near the umbones while the outer calcareous layer is thin and contains a high proportion of organic material. The shell conchiolin consists mainly of protein, varying in composition, but much of it strengthcned by quinone-tanning, and in ccrtain regions probably by the presence of appreciable quantities of chitin. The ligament, although superficially resembling an amphidetic structure, is opisthodetic, the extcnsion anterior to the umbones consisting of anterior outer layer only.
The mantle is characterized by an extension of the outer fold of the mantle margin which has effected equally both the inner and outer surfaces of this fold. The secretory epithelium and the modified pallial musculature, contraction of which results in the intucking and plaiting of the periostracum, is dcscribed. Simple tubular oil glands open at the mantlo margin and are responsible for the water-repellent nature of the periostracum.
The form of the mantlelshell and that of the enclosed body are discussed and compared with those of other bivalves in which elongation of the mantle/shell is achieved in a different way. It is concluded that the mantlelshell of Solemya is of little value in determining its relationships, and that the greatly elongatod ligament, the edentulous hinge and the flexible shell are all adaptations to a specialized mode of life.     

DISTRIBUTION AND PRODUCTION OF THREE HYDROBIA SPECIES (GASTROPODA: HYDROBIIDAE) IN A SHALLOW COASTAL LAGOON IN THE BAY OF CADIZ, SPAIN

The abundance, life span, growth and production of the mud snailsHydrobia minoricensis, H. ulvae and H. ventrosa in a semi-naturallagoon system were studied by taking monthly samples at threesites during 1991 and 1992. The most abundant species, H. minoricensisoccurred at mean densities of 12834 to 26264 snails m^–2(10.7 to 25.8g dry weigh m^–2), depending on the site.The least abundant species, H. ulvae, occurred at mean densitiesof 185 to 353 snails m^–2 (3.2 to 2.2g dry weight m^–2).The numerical abundance and biomass of the three Hydrobia specieswere positively related to the biomass of benthic macroalgae(P<0.01). Although H. ulvae egg capsules were recorded throughoutthe year, newly hatched snailsof this species were not observed,in contrast to the other two species. The early spring and summercohorts of H. minoricensis and H. ventrosa seemed to be themost numerous. The average life spans of these two species wereestimated to be about 18 and 13 months respectively. Annualproduction estimates for the whole lagoon system were 29.0 (6.3),5.5 (0.8) and 5.2 (1.0)g dry weight (ash-free dry weight) m^–2yr^–1 for H. minoricensis, H. ulvae and H. ventrosa respectively.The annual P/B ratio was about 2 for H. minoricensis and H.ventrosa. (Received 5 July 1994; accepted 5 October 1994)     

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;     

INTER-YEAR CONTRAST IN NUMBERS AND ANNUAL PRODUCTION OF THE CEPHALASPIDEAN OPISTHOBRANCH RETUSA OBTUSA (MONTAGU) AND THE POSSIBLE INFLUENCE OF WIND-SPEED

Retusa obtusa was far more abundant throughout August 1986-July1987 than in all 12 corresponding months of April 1992-March1993. The greatest difference was when the highest of all counts(3232 m^-2) in May 1987 was 18-times that in May 1992. The leastdifference was a 2.4-fold discrepancy between the two Septembercounts. Mean monthly counts for the 2 periods were 781.4 m^-2(1986-87) and 123.6 (1992-93): a difference of 6.3-times. Biomasssimilarly differed between the 2 periods by factors between2.5 (July) and 12 (March), with a 5.3-fold difference betweenmean monthly biomass values. Annual production was accordinglyestimated as 618.3 (1986-87) and 152.4 (1992-93) mg AFDW m^-2y^-1; so differing by a factor of 4. Yet, in both these 1-yearperiods of observation, the general pattern of annual changewas the same, with settlement in February-May, growth from springto winter, most spawning in December-March and death of adultsat about 1 year in April-May. It is likely that the winter-springperiod of early development in benthic eggs followed by pelagicdispersal, and then settlement in February -May, is a periodof especial vulnerability. In particular, it seems likely thatduring January-May, markedly higher wind-speeds in 1992 thanin 1987 might have contributed to the huge difference in numbersof recruits (Received 2 February 1994; accepted 18 March 1994)     

Direct innervation of the mantle edge gland by neurosecretory axons in Helisoma duryi (Mollusca: Pulmonata)

Summary The mantle edge gland of Helisoma duryi is innervated by neurosecretory axons from the pallial nerves. Synaptoid contacts occur between axons and gland cells, and there is ultrastructural evidence for the release of neurosecretory material. The mantle edge gland contributes to the deposition of periostracum during shell formation, and direct neurosecretory innervation may control shell growth and regeneration.Supported by a National Research Council of Canada Grant (A-4673) and Negotiated Grant D-61