Mechanism of an epibiont burden: Crepidula fornicata increases byssus thread production by Mytilus edulis

Epibionts of mussels can have detrimental effects on their basibionts,such as reduced growth rates, lower fecundity, increased mortalityand an enhanced risk of dislodgement of the overgrown bivalvesdue to stronger hydrodynamic forces. In blue mussels Mytilusedulis, the epibiotic American slipper limpet Crepidula fornicatareduces growth and survival. In a field experiment we testedthe hypothesis that an enhanced byssus thread production withhigh energetic costs for the mussels due slipper limpet epibiontsis the underlying mechanism for the epibiont burden. Byssusthread production in overgrown mussels was twice as high asin unfouled M. edulis (11 ± 0.9 and 5.4 ± 0.6byssus threads/mussel/day, respectively). A control experimentrevealed intermediate byssus thread production (8.4 ±0.8 byssus threads/mussel/day) in mussels cleaned of C. fornicataat the beginning of the experiments, indicating that C. fornicatais responsible for the effects observed. We conclude that increasedbyssus production in fouled M. edulis is a functional responseto higher drag caused by epibionts and that it is associatedwith increased energy expenditure that reduces allocation ofresources for other processes such as growth, reproduction andsurvival. Such indirect effects of epibionts, mediated by anenhanced byssus production, may be widespread in byssus-producingbivalves, with important implications for their population dynamics. (Received 12 January 2006; accepted 21 November 2006)     

Specialized cilia of the byssus attachment plaque forming region in Mytilus californianus

The distal depression of the ventral pedal groove of Mytilus californianus was investigated by scanning and transmission electron microscopy. This part of the byssus forming system is responsible for the formation of the attachment plaque of the byssus thread. The longitudinal pedal ducts open into this area and the floor of the distal depression is covered by specialized cilia which terminate as biconcave flattened discs or “paddles.” The disc is formed by a 360° curvature of the axoneme tip within the ciliary membrane. The diameter of the disc is about 1.33 μ while that of the shaft portion is 0.24 μ. There are about 11 cilia per square micron of surface area and the necks of the cilia are separated from each other by a web-like extension of apical cytoplasm extending from the epithelial cells. It is proposed that these specialized cilia function as microscopic spatulas for the application of the adhesive plaque material to substrate surfaces. The pattern of surface convection currents seen in vivo tends to support this hypothesis.     

MECHANICAL PROPERTIES OF MUSSEL BYSSUS THREADS

The byssus threads of the common mussel, Mytilus edulis L.,have been tested mechanically and the results from the testsrelated to the ecology of the animal. The threads are mechanicallysimilar to other crystalline polymers such as polyethylene havinga modulus of about 10⁸N m^–2 and a long relaxation time.Resilience of 60% is similar to tendon; ultimate strain is aboutfive times that of tendon at 0.44. The thread is laid down witha prestrain of 10% and so guys the mussel in position. Calculationshows that a mussel with 50 byssus threads would be able toresist all but severe winter storms (Received 1 December 1978;     

THE FUNCTIONAL ANATOMY OF THE NERITACEAN LIMPET PHENACOLEPAS OMANENSIS BIGGS AND SOME COMPARISON WITH SEPTARIA

The shell of Phenacolepas and Seplaria is produced by a disproportionateexpansion of the last whorl of the typical neritid shell, withthe right and left columellar muscle an extension of that partarising from the shell and not the columella. The columellais a posterior ridge near the level of the protoconch. In contrastto zeugobranchs the mantle cavity is large, extending beneaththe viscera: the organization and disposition of the organsrelated to it are similar to those of Nerita. Poor development of pedal musculature of Phenacolepas may berelated to the sheltered environment. As in other limpets thereis a marginal pedal flange acting as a seal around the areaof low pressure which arises when the foot is used as a sucker.The internal anatomy reflects neritacean conservatism, but thereare characters, previously undescribed, which justify the separationof the genus in a separate family, the Phenacolepadidae: theelaboration of the mantle edge to resemble that of some zeugobranchs;the retention of an internal operculum; relatively sparse developmentof pedal musculature; the loss of the right auricle; erythrocytesin the blood; a vaginal papilla related to an ovipositor; theabsence of a crystal sac and modifications of the nervous systemto give more direct nervous pathways. (Received 12 September 1982;     

Byssus thread strength in the mussel, Mytilus edulis

Mytilus edulis attaches to the substratum by means of a proteinaceous byssus complex. This consists of three portions: a root, embedded in the pedal tissues, a stem, continuous with the root but external to the body and a number of byssus threads attached proximally to the stem and distally to the substratum via adhesive discs. Byssus strength varies seasonally on the shore, in response to changes in wave action (Price, in press). As a decline in byssal attachment strength implies a decline in strength of the constituent threads, a study was undertaken to establish the extent to which byssus thread strength is determined by age. The ultimate tensile stress, ultimate tensile strain and Young's Modulus were measured in threads of known age and length and a stepped regression performed on the results. It was found that age and length correlate significantly with tensile stress and Young's Modulus. Length is a less important influence than age on tensile stress but has a greater effect than age on Young's Modulus. Tensile strain is independent of both length and age.     

The ecology and life cycle of a population of Modiolarca tumida (Hanley, 1843) (Bivalvia: Mytilidae) of the coast off north-eastern England

The life history of a population of Modiolarca tumida occurringoff the Northumberland coast is described. Modiolarca tumidalives primarily in the tests of ascidians. On settlement theyoung spat seek an ascidian host, climb close to the inhalentand exhalent apertures of the host and there wrap the test aroundthemselves using the byssus. Initially, there is a preferencefor Ascidiella scabra as the host species but, with increasingsize, Modiolarca is more and more often found in Polycarpa pomaria.While numbers per ascidian vary, usually <5 occur in eachhost. However, immediately following settlement very occasionally100+ have been recorded. Modiolarca tumida is relatively short-lived,with few living longer than two years. Spawning occurs at theend of July and August at the end of its first year of lifeand at the same time in subsequent years in the case of theminority that survive. (Received 13 February 2007; accepted 9 November 2007)     

PEDAL AND OPERCULAR SECRETORY GLANDS OF POMATIAS, BITHYNIA AND LITTORINA

The histochemical properties of the pedal and opercular glandcells of three prosobranchs from different habitats were examined.The suprapedal gland of Pomatias elegans contained 3 gland celltypes producing mucoprotein, protein and sulphated muco-polysaccharide.The ventral surfaces of the foot were devoid of gland cellsexcept in the median furrow in which two cell types producea neutral and a sulphated mucopolysacharide. The dorsal surfaceof the foot possesses 5 cell types which produce a variety ofmucosubstances. The anterior pedal gland of Bithynia tentaculata produces neutraland weakly acidic mucoprotein from one cell type. The ventralsurface of the foot is generally populated by gland cells oftwo types producing acid mucopolysaccharide and protein. A furthercell type producing carboxylated mucopolysaccharide is restrictedto a transverse band mid-way down the foot. The dorsal surfaceof the foot is covered by mucus from three cell types producinga variety of mucosubstances. The anterior pedal gland of Littorina littorea possesses twocell types both of which secrete mucoprotein. The ventral surfaceof the foot secretes sulphated and carboxylated mucopolysaccharideand a mucoprotein from three cell types which form a thick subepi-dermalglandular layer. The dorsal surface is lubricated by a sulphatedmucopolysaccharide and a mucoprotein. The secretory cells of the dorsal surface extend into the operculargroove and disc whose specialised cells generally secrete mucopolysaccharideand at least two differently staining proteins. ^*Present address: Department of Zoology, University of Durham,Durham City, U.K. (Received 19 August 1986;     

Neuroethology of Melibe leonina Swimming Behavior

The nudibranch Melibe leonina swims by rhythmically flexingits body from side to side at a frequency of 1 cycle every 2–5sec. Melibe swim spontaneously, when they are dislodged fromthe substrate, or when they come in contact with predatory seastars,such as Pycnopodia helianthoides. Intracellular recordings obtainedfrom semi-intact swimming Melibe reveal a population of 15 swimmotoneurons (SMNs) in each pedal ganglion. In general, SMNsin one pedal ganglion fire out-of-phase with SMNs in the oppositepedal ganglion, resulting in rhythmic side-to-side bending movements.In isolated brains, recordings from SMNs yield similar results,indicating the existence of a swim central pattern generator(CPG). There is no evidence for synaptic interactions betweenSMNs and either inhibiting or exciting SMNs has no impact onthe swim pattern. The SMNs are driven by a CPG consisting of4 interneurons; 2 in the cerebropleural ganglia and 1 in eachpedal ganglion. Appropriate bursting activity in the swim interneuronsis necessary for swimming to occur. Either hyperpolarizationor depolarization of any of the 4 CPG interneurons disruptsthe normal swim pattern. Swimming behavior, and the fictiveswim motor program expressed by the isolated brain, are inhibitedby light and nitric oxide donors. NADPH-diaphorase stainingand nitric oxide synthase (NOS) immunocytochemistry of Melibebrains suggests the source of nitric oxide might be a pair ofbilaterally symmetrical cells located in the cerebropleuralganglia.     

The effect of food availability on byssogenesis by the zebra mussel (Dreissena polymorpha Pallas)

The effect of varying algal availability on byssal thread productionby re–attaching zebra mussels (Dreissena polymorpha) wasquantified. The byssal apparatus was severed and mussels allowedto re–attach to a hard substratum for a 21 day periodduring which they were fed at algal concentrations of 0.0, 0.1,0.5, 1.0 or 2.0 mg C l^-1. Byssal thread production was quantifiedby counting the number of new attachment plaques present eachday. Results showed that starved mussels continued to partition organiccarbon towards thread production but the resultant byssal masswas compromised, containing fewer threads than those producedby fed mussels. The daily average byssal thread production bymussels fed at 2.0 mg C l^-1 was greater than that of starved musselsand the final mean dry soft tissue weight higher. At algal Cconcentrations below maintenance requirements byssal threadproduction was elevated compared to starved mussels, but therewas no concurrent increase in soft tissue. This suggests thatbelow maintenance levels assimilated carbon was solely partitionedtowards byssus production and stored reserves may have beenutilized. The ratio of organic carbon contained in the byssusto that in the soft tissues remained relatively constant acrossall feeding levels. This suggests that the carbon content ofthe byssus is a constant function of that of the soft tissuemass. These results may explain seasonal variation in attachmentstrength of numerous byssate species and seasonal vertical migrationsby D. polymorpha. Present Address-Queen Mary & Westfield College, Dept of Biology,University of London, Mile End Road, London E1 4NS, UK (Received 16 March 1998; accepted 30 September 1998)     

盐度对企鹅珍珠贝足丝分泌的影响

企鹅珍珠贝（Pteria penguin）是生产附壳珍珠的大型海水经济贝类,其依靠强壮的足丝将自身固定在硬质基底上,抵抗水流的冲击和抵御被捕食等。足丝分泌和足丝的形状很容易受到环境的影响,本实验采用盐度30为低盐度组、盐度35为中盐度组和盐度40为高盐度组,研究这3种盐度对企鹅珍珠贝足丝分泌、足丝直径和足丝拉力的影响,通过单因素方差分析法（LSD法）分析这三个足丝相关指标在3种盐度组间是否存在显著性差异。结果显示,3种盐度下企鹅珍珠贝足丝附着率无显著差异,但在整个实验周期72 h内,中盐度组的足丝分泌总数为（48.7 ± 15.1）根,显著高于低盐度组的（24.7 ± 5.0）根和高盐度组的（13.3 ± 1.5）根。在实验的前6 h内,中盐度组的足丝首次附着率显著高于低盐度组和高盐度组（P < 0.05）,但在后续的12 h、18 h、30 h、42 h、54 h和66 h这6个时间点,3个盐度组的足丝首次附着率均无显著性差异。足丝直径未受盐度变化的影响,但盐度对足丝拉力具有显著影响,中盐度组的足丝拉力显著高于低盐度组和高盐度组（P < 0.05）。上述结果表明,企鹅珍珠贝为适应一定范围内盐度的改变,会在短时间内通过抑制足丝分泌来减少能量消耗,随着对环境的适应足丝分泌会恢复。盐度影响足丝分泌且对足丝拉力影响显著,但对足丝直径无明显影响。本研究可以为企鹅珍珠贝养殖及珍珠插核培育提供理论基础。     

BIOCHEMICAL COMPOSITION OF HELIX SNAILS: INFLUENCE OF GENETIC AND PHYSIOLOGICAL FACTORS

This paper describes the biochemical composition of differentspecies (Helix lucorum, Helix pomatia) and sub-species of snails(Helix aspersa aspersa, Helix aspersa maxima) reared in thesame conditions with a feed (‘Helixal’) speciallydesigned for edible snails. In addition, the composition ofwild H. pomatia and H. lucorum is presented to allow comparisonbetween snails of different origins. Analyses determined thepercentages of proteins, lipids and minerals. They reveal bothsimilarities and differences in composition according to thespecies and the part analysed (whole body, pedal mass, and visceralmass). H. pomatia contains the highest percentage of mineralmatter and the lowest percentage of lipids. Surprisingly, proteincontents are slightly different between artificially rearedH. aspersa maxima of 3 months old and wild H. pomatia. The resultsmake it possible to evaluate nutritional quality of snails withthe composition of the body of four edible snail species. (Received 16 September 1996; accepted 24 May 1997)     

A variety of Mytilus inhibitory peptides in the ABRM of Mytilus edulis: isolation and characterization.

1. Five species of Mytilus inhibitory peptides, MIP1-5, were isolated from acetone extracts of the anterior byssus retractor muscle (ABRM) of Mytilus edulis. MIP1 and MIP2 were shown to be S2-MIP and A2-MIP, respectively, first isolated from the pedal ganglia of the animal. 2. All the five peptides had a common C-terminal structure of -Pro-Xaa-Phe-Val-NH2, which was shown to be important for their biological activity. 3. The five MIPs showed similar inhibitory effects on contractions of the ABRM but did not affect catch tension and its relaxation. 4. In addition to the MIPs, catch-relaxing peptide (CARP) was also found in the ABRM.     

Byssus production ability and degree of byssal-gland development in the infaunal clam Ruditapes philippinarum

Byssus production of Ruditapes philippinarum clams becomes reduced with growth. This tendency is well recognized but has not been analysed in detail. Additionally, it remains uninvestigated whether the lack of competence to produce byssus threads in the adult stage is caused by atrophy of the byssal glands or not. The objective of this study was to evaluate the byssus production ability of clams through the juvenile to adult stages and to examine the importance of two endogenous factors (i.e. shell size, somatic condition) in determining the byssus production probability (proportion of clams with byssus production in a population). This study also histologically confirmed the presence of byssal glands in juvenile to adult clams. For these purposes, field surveys to investigate the relationship among byssus production, shell size and somatic condition of clams collected from four intertidal sites and a histological study for byssal glands of the clams was conducted. This study revealed that byssus production probability decreases with increasing shell size and declining somatic condition and that the lack of byssus production is not caused by the loss of the byssal glands.     

Anatomy of Bathyacmaea secunda Okutani, Fujikura & Sasaki, 1993 (Patellogastropoda: Acmaeidae)

Anatomy of a vent-endemic patellogastropod limpet, Bathyacmaeasecunda, was examined by gross dissection and serial sections.It was revealed that B. secunda is anatomically distinct fromother patellogastropod taxa in that (1) the intestine runs throughthe ventricle; (2) the ventral approximator muscle of odontophoralcartilages is ventrally single-layered; (3) there is a pairof radular teeth with long shafts; and (4) the statocysts areisolated from pleural and pedal ganglia. In addition, the speciesis characterized by short pallial margin papillae, lack of pallialstreaks, presence of a ctenidium, obliquely tubular salivaryglands, simple gut configuration and acmaeoidean type of buccalmass musculature and odontophoral cartilages. The comparisonin anatomical and shell microstructural characters suggeststhat B. secunda shares no unique similarities with the Patelloideaor Neolepetopsidae and, therefore, is not closely related tothese groups. In contrast, a number of similarities were foundbetween B. secunda and acmaeid species. These results supportthe current systematic position of Bathyacmaea within the Acmaeidae,although anatomical data from some related genera are stillinsufficient. (Received 3 September 2005; accepted 25 January 2006)     

THE INNERVATION AND PHYSIOLOGY OF THE EXTRINSIC BUCCAL RETRACTOR MUSCLES OF PHILINE APERTA (LINNAEUS)

Two buccal mass retractor muscles of Philine are innervatedby at least 4 excitatory motoneurons, whose cell bodies liein the buccal and the cerebral ganglia. The muscle fibres respondto action potentials generated in the motoneurons or their axonswith excitatory junction potentials (ejps), each of which isfollowed by a small twitch-like contraction. Both the electricaland mechanical responses facilitate and summate with repetitivestimulation. A large ventrally located cerebral neuron (VGC) inhibits tensiondevelopment in the muscle by reducing the amplitude of the excitatoryjunction potentials from and identified buccal motoneuron. Acetylcholinereversibly depolarises and causes tonic contraction of the muscles.This action is partially antagonised by hexamethonium, whichalso blocks the ejps from two axons in the buccal and one inthe pedal nerve 9. 5-Hydroxytryptamine potentiates the ejp fromthe identified buccal motoneuron and enhances the rate of relaxation.Histamine reduces the amplitude of the presumed cholinergicbuccal nerve ejps, but does not affect the hexamethonium sensitiveejp in the pedal nerve 9. In this respect its action resemblesthat of the ventral giant cell.     

Functional Morphology of Perihelia conradi Relative to Shell-Penetration

The pholad, Penitella conradi, is found along the Californiacoast in the calcareous shell of the abalone, Haliotis rufescens.These pholads penetrate the abalone shell, and when they breakthrough the inside of the shell they cease boring, secrete acallum, and then become sexually mature. The normal adult isa stenomorphic form,defined by Bartsch as an animal whose sexualmaturity is induced by over-crowding or insufficient substratumin which to bore. In the case of P. conradi, sexual maturityis always induced by the spatial limits of the substratum, thatis, the relatively thin abalone shell. The role of mechanical abrasion by the valves of P. conradiis minor. Experiments indicate that the teeth of P. conradiare worn at a greater rate than the polished shell of the abalone. The boring process in P. conradi proceeds mainly by chemicaldissolution of the calcareous substrate. The pathway of thesolvents is unknown. It may be through the organic matrix, orthe solvent may react directly with the crystals. Mechanicalabrasion helps to remove loosened crystals and/or organic matrixwhich are then carried to the exterior by the ciliary currentsflowing in through the pedal gape and out through the exhalentsiphon.     

Byssus of the Green-Lipped Mussel <Emphasis Type="Italic">Perna viridis</Emphasis> (Linnaeus) as a Biomonitoring Material for Zn

Recently, Yap et al. (2003) suggested that the byssus of the green-lipped mussel Perna viridis can be a biomonitoring material for Zn although further validation is required. In this work, we did a simple correlation study between Zn concentrations in the byssus (and soft tissue) and in different geochemical fractions of the sediment. A significant (P < 0.01) Pearsons correlation coefficient (R = 0.84) between the Zn concentrations in the byssus and soft tissue indicated that the Zn level in the byssus is highly correlated to its level in the soft tissue and that the byssus could act as an excretion route for Zn. Higher R-values were found between the byssus-easily or freely, leachable and exchangeable, byssus-acid-reducible, byssus-oxidizable-organic and byssus-nonresistant fractions of the sediment, and the byssus-Zn concentration in the total sediment when compared to those found between the soft tissue and the same geochemical fractions. This indicated that the byssus was more reflective of Zn contamination in the field environment than the soft tissue. Therefore, the data further support the use of the byssus as a biomonitoring material for Zn as was originally suggested by Yap et al. [3].Original English Text Copyright © 2005 by Biologiya Morya, Yap, Ismail, Tan.This article was submitted by the authors in English.     

A variety of Mytilus inhibitory peptides in the abrm of Mytilus edulis: Isolation and characterization

1. Five species of Mytilus inhibitory peptides, MIP_1–5, were isolated from acetone extracts of the anterior byssus retractor muscle (ABRM) of Mytilus edulis. MIP₁ and MIP₂ were shown to be S²-MIP and A²-MIP, respectively, first isolated from the pedal ganglia of the animal.2. All the five peptides had a common C-terminal structure of -Pro-Xaa-Phe-Val-NH₂, which was shown to be important for their biological activity.3. The five MIPs showed similar inhibitory effects on contractions of the ABRM but did not affect catch tension and its relaxation.4. In addition to the MIPs, catch-relaxing peptide (CARP) was also found in the ABRM.     

Fission in sea anemones: integrative studies of life cycle evolution

Sea anemones (Phylum Cnidaria; Class Anthozoa, Order Actiniaria)exhibit a diversity of developmental patterns that include cloningby fission. Because natural histories of clonal and aclonalsea anemones are quite different, the gain and loss of fissionis an important feature of actiniarian lineages. We have usedmitochondrial DNA and nuclear intron DNA phylogenies to investigatethe evolution of longitudinal fission in sixteen species inthe genus Anthopleura, and reconstructed an aclonal ancestorthat has given rise at least four times to clonal descendents.For A. elegantissima from the northeastern Pacific Ocean, atransition to clonality by fission was associated with an up-shorehabitat shift, supporting prior hypotheses that clonal growthis an adaptation to the upper shore. Fission in Actiniaria likelyprecedes its advent in Anthopleura, and its repeated loss andgain is perplexing. Field studies of the acontiate sea anemoneAiptasia californica provided insight to the mechanisms thatregulate fission: subtidal Aiptasia responded to experimentallydestabilized substrata by increasing rates of pedal laceration.We put forth a general hypothesis for actiniarian fission inwhich sustained tissue stretch (a consequence of substratuminstability or intrinsic behavior) induces tissue degradation,which in turn induces regeneration. The gain and loss of fissionin Anthopleura lineages may only require the gain and loss ofsome form of stretching behavior. In this view, tissue stretchinitiates a cascade of developmental events without requiringcomplex gene regulatory linkages.