Effects of temperature,salinity and agitation on byssus thread formation of zebra musselDreissena polymorpha

Byssus thread production ofD. polymorpha under different conditions of temperature, salinity and agitation were studied in the laboratory. The acclimation to salinity and temperature greatly affects the byssus production ofD. polymorpha. Byssus production of mussels was significantly reduced when temperature increased beyond 20°C and decreased below 10°C. Mussels with cut threads (for counting), produced a substantially increased number of threads. However, mussels with uncut byssus threads were comparatively more mobile. Byssus production of mussels did not vary significantly at salinities up to 3. Beyond this salinity byssus production was reduced significantly. Mussels increased their byssus production with increasing frequency of agitation.     

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)     

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)     

Citrate synthase from Antarctic ciliates: adaptation to low temperatures and comparison with temperate ciliates

The activity of citrate synthase (CS), a mitochondrial enzyme in the energy metabolism pathway, was studied in Euplotes focardii (Hypotrichia) and Uronema marinum (Scuticociliatia), isolated from samples of Antarctic seawater and, for comparison in E. vannus and U. nigricans, related ciliates from the Adriatic. The temperature-dependent characteristics of CS were investigated in the range 0–55°C, to evaluate mechanisms of thermal regulation, such as optimal temperature levels, activation energy, and kinetic parameters. CS activity was up to double that recorded in temperate organisms at reaction temperatures between 0 and 10°C. The optimal temperature for enzymatic activity was independent of Tamb. Activation energy for CS was significantly lower in Antarctic ciliates than in temperate ones, indicating a significant increase in the reaction rate. KM at various temperatures of the reaction substrates indicated the higher affinity of CS for acetyl-CoA in both Antarctic organisms at lower temperatures. The data for thermal profiles for KM, showing greater differences between E. focardii and E. vannus than between U. marinum and U. nigricans, support the distinctive physiological characteristics of each species. E. focardii is an endemic and psychrophilic species, whereas U. marinum is a typically ubiquitous species, more adaptable to environmental changes.     

Byssogenesis in the juvenile pink heelsplitter mussel,Potamilus alatus (Bivalvia: Unionidae)

The North American pink heelsplitter (Potamilus alatus) differs from most freshwater mussels in China by the ability to secrete an ephemeral byssus during its juvenile stage. In the present study, light microscopy, scanning electron microscopy (SEM), and transmission electron microscopy (TEM) were used to investigate this ephemeral byssal structure, and amino acid composition was analyzed and compared with that of other species. The results revealed that the byssus consists of a long byssal thread and a few adhesive plaques which are randomly set up along the thread and assembled by petioles. There is a thin but distinctive cuticle with a continuous homogeneous matrix surrounding the byssal thread. Structural variation occurred when the byssal thread was differentially stretched. Four‐stranded helical primary fasciculi, which form a stable rope‐like structure, become evident after removal of the cuticle. The primary fasciculi consist of bundles of hundreds of parallel secondary fasciculi, each measuring about 5 μm in diameter. All evidence indicates that the byssus of the pink heelsplitter has a significantly different macrostructure and microstructure than the permanent byssus of the marine mussel Mytilus. Byssogenesis ceases when juveniles exceed 30 mm in length, although it varies greatly even among juveniles of similar size. Byssus formation is influenced by substrate type. The unique characteristics of the byssus have important advantages for survival, transition, and aggregation during the early life history. This study not only provides first insight into the structure of the ephemeral byssus and its relationship to freshwater mussel development and growth, but also suggests possibilities for the synthesis of novel biopolymer materials particularly useful in freshwater ecosystems. J. Morphol. 276:1273–1282, 2015. © 2015 Wiley Periodicals, Inc.     

A difference in optomotor behaviour of two Antarctic nototheniid fishes is correlated with the presence of a choroid<Emphasis Type="Italic"> rete mirabile</Emphasis> and Root effect

The hypothesis that fish vision is influenced by the presence of a choroid rete mirabile (an oxygen-concentrating apparatus in the eye) and the Root effect was tested using the optomotor response in two nototheniid species from Antarctica. Pagothenia borchgrevinki, an active cryopelagic zooplanktivore, does not exhibit a major Root effect and the rete is absent. In contrast, the sluggish, benthic nototheniid, Trematomus bernacchii, possesses both a rete and a significant Root effect. Whole-eye ocular PO2 was higher in T. bernacchii (18.6 kPa) than in P. borchgrevinki (18.1 kPa) and the optomotor response of T. bernacchii was positively influenced by subtended angle on a moving background whereas P. borchgrevinki did not respond to any subtended angle (0–600 min of arc). The optomotor response of T. bernacchii may therefore have been facilitated by elevated ocular PO2 due to the presence of the choroid rete and the Root effect.     

Photoprotection in evergreen Mediterranean plants during sudden periods of intense cold weather

The photoprotective responses to an abrupt period of exceptional cold weather were studied in several Mediterranean evergreen species with different ecological requirements. The same pattern of response was observed in most of the species with little change in hydrophilic antioxidants (ascorbate and glutathione) and the carotenoid pool, an increase in the content of α -tocopherol, and a night retention of de-epoxidised xanthophylls (antheraxanthin and zeaxanthin). The accumulation of these xanthophylls correlated with a sustained decrease in maximal photochemical efficiency (Fv/Fm). This reduction in the rate of electron transport would reduce the production of superoxide in photosystem I, as well as the subsequent hydrogen peroxide and hydroxyl radical. Thereby if any transitory photooxidative stress is produced under these conditions it should be due mainly to the formation of singlet oxygen by triplet excited chlorophyll within the antenna. Since α-tocopherol is the main scavenger of singlet oxygen and lipid peroxy radicals, the large increase of this antioxidant within the species could be enough to compensate for the higher degree of photooxidative stress, playing an essential role in the survival of vegetation during the incidence of exceptional cold fronts in the Mediterranean region.     

Bacillus naphthovorans sp. nov. from oil-contaminated tropical marine sediments and its role in naphthalene biodegradation

. A Bacillus sp., designated as strain MN-003, was isolated as the dominant cultivatable naphthalene-degrading organism from oil-contaminated tropical marine sediments. Strain MN-003 is strictly aerobic, rod-shaped, Gram-positive, catalase positive, oxidase negative, and forms endospores. Strain MN-003 grew at salinities ranging from 0.28 to 7.00% and temperatures ranging from 15 to 41°C. Phylogenetic analyses reveal that strain MN-003 is most similar to Bacillus sp. VAN14, with a 16S rRNA sequence identity of 97.9%. Based on taxonomic and 16S rRNA data, strain MN-003 was named Bacillus naphthovorans sp. nov. When grown with naphthalene as sole carbon source, strain MN-003 had a maximal specific growth rate (µmax) of 0.32ǂ.03 h^–1, and a half-saturation constant (Ks) of 22.3dž.2 µM. A batch study of the tropical marine sediments enriched with naphthalene showed that cells of the Bacillus genus grew to become dominant members of the microbial community. The bacilli comprised 39.5Lj.5% of the microbial fraction after 20 days of enrichment.     

Microsatellite Polymorphism and the Population Structure of the Black Tiger Shrimp (Penaeus monodon) in Thailand

: Genetic variation and differentiation of Thai Penaeus monodon from five geographic locations (Chumphon, Trad, Phangnga, Satun, and Trang) were investigated using five microsatellite loci (CUPmo18, Di25, Di27, CSCUPmo1, and CSCUPmo2). The number of alleles across the five loci ranged from 19 to 30, and heterozygosities ranged from 0.49 to 0.95. The mean number of alleles and effective number of alleles per locus were 21.0 to 26.6 and 13.1 to 20.4, respectively. The average heterozygosity across all investigated samples was 0.78, indicating high genetic diversity in this species. Geographic heterogeneity analysis of the results from two of the loci, CUPmo18 and Di25, showed significant differences among the Gulf of Thailand (Trad and Chumphon) but not the Andaman samples. Comparison between regions revealed significant heterogeneity of the Andaman and Trad P. monodon (P < .001), whereas those from Chumphon and the Andaman were genetically similar (P > .05). Significant genetic differentiation was consistently observed between the Andaman-Trad samples (FST = 0.0101, P < .0001) and the Chumphon-Trad samples (FST = 0.0101, P < .0001). On the basis of our analyses, the investigated samples from five geographic locations were allocated to three distinct populations composed of the Andaman Sea (A), Chumphon (B), and Trad (C).     

Effects of artificial epibionts on byssogenesis,attachment strength,and movement in two size classes of the blue mussel,Mytilus edulis

Blue mussels (Mytilus edulis) can alter the strength of byssal attachment and move between and within mussel aggregations on wave‐swept shores, but this movement ability may be limited by epibiont fouling. We quantified the effects of artificial epibiont fouling on the production of byssal threads, attachment strength, and movement in two size classes of blue mussels. In a factorial experiment, large epibiont‐covered mussels produced more functional byssal threads (i.e., those continuous from animal to substrate) after 24 h than large unfouled and small fouled mussels, but not more than small unfouled mussels. Small unfouled mussels formed and released more byssus bundles compared to any other treatment group, which indicates increased movement. Conversely, epibiont fouling resulted in decreased numbers of byssus bundles shed, and therefore reduced movement in small mussels. Epibiont‐covered mussels started producing byssal threads sooner than unfouled mussels, while small mussels began producing byssal threads earlier compared to large mussels. Mean attachment strength from both size classes increased by 9.5% when mussels were artificially fouled, and large mussels had a 34% stronger attachment compared to small mussels. On the other hand, a 2.3% decrease in attachment strength was found with increasing byssus bundles shed. Our results suggest that fouling by artificial epibionts influences byssal thread production and attachment strength in large mussels, whereas epibionts on small mussels impact their ability to move. Mussels are able to respond rapidly to fouling, which carries implications for the dynamics of mussel beds in their intertidal and subtidal habitats, especially in relation to movement of mussels within and among aggregations.     

Location and analysis of byssal structural proteins of Mytilus edulis

The acellular attachment organ (byssus) of the marine mussel Mytilus edulis L. is composed of threads that emanate from the body of the mussel to adhesive discs that anchor the threads to rocks, sand and other mussels. Three proteins have been purified by immunohistological methods and located to specific regions of the byssus. A collagenous protein with subunit molecular weights of 53,000, 55,000 and 65,000 is found in the matrix of the elastic thread region. Its 73,000-MW precursor was extracted from foot glands in the area proximal to the animal body and was identified by immune cross-reactivity. A cystine-rich, acidic protein was found in all regions of the byssus associated with a third protein, the polyphenolic protein. The L-dopa-containing polyphenolic protein appears in the cortex of the entire thread and adhesive plaque and at the substrate-plaque interface. Antiserum to this protein stains spherical vesicles in the phenol gland of the foot. Using immuno-electrophoretic methods, the polyphenolic protein and the cystine-rich protein were shown to form high molecular weight aggregates with aging of the byssus.     

Marine mussels Brachidontes variabilis selected smaller places of refuge and enhanced byssus production upon exposure to conspecific and heterospecific cues

Refuge selection and byssus production as anti-predator responses were examined in mussels Brachidontes variabilis exposed to cues either released from damaged or intact conspecifics, from damaged or intact heterospecific mussels Perna viridis sympatric with B. variabilis or from shrimp meat as novel cues. Most of the mussels from all treatment groups would seek and stay in a refuge for the first few hours, with a significantly higher number of mussels from the damaged conspecific group seeking refuge. More mussels preferred a smaller refuge when they were exposed to conspecific or heterospecific cues; mussels from other treatments did not select particular sizes of refuge. Damaged conspecific and heterospecific cues elicited the greatest responses in byssus production with more byssal threads being produced, which were also longer and thicker. Novel cues elicited medium levels of response, suggesting that some common cues from injured or dead individuals from different taxa were released, which induced anti-predator responses in B. variabilis.     

Morphological Adaptations of the Mussel Crenomitilus grayanus (Bivalvia) to Attached Life

The morphology of the mussel Crenomytilus grayanus from different biotopes manifesting the adaptation of this mollusk to attached life on the bottom surface was studied. It was established that colonization of rocky coastal areas with active hydrodynamics by mussels is related to retardation of linear growth rates, the formation of convex shells, and vigorous development of retractor muscles of the foot and byssus. An environment protected from the effect of surfs on silted bottoms facilitated the fast growth of mussels, which resulted in the formation of an elegant flat shell. In that biotope mussels are specified by a rather poor development of the muscles responsible for its attachment to the substrate.     

Primary production,light absorption and quantum yields of phytoplankton from the Bellingshausen and Amundsen Seas (Antarctica)

Chlorophyll concentration, light intensity, primary production, light absorption and quantum yield were measured between 12 January 1994 and 27 March 1994 in the Bellingshausen and Amundsen Seas. Primary production and quantum yield within Bellingshausen and Amundsen Seas were typical of the high-nutrient, low-chlorophyll area of the Southern Ocean while small variations were found as a result of local conditions. Chlorophyll a (chl a) concentrations were generally low (<1 μg l⁻¹) in the water column, while in cases of blooms it reached 7–8 μg l⁻¹. The light intensity at which photosynthesis approaches saturation varied between 59 and 105 μmol q m⁻² s⁻¹.The initial slope of the photosynthesis curve varied between 0.02 and 0.07 μg C (μg chl a)⁻¹ h⁻¹ (μmol q m⁻² s⁻¹)⁻¹. The maximal photosynthetic rate at light saturation ranged between 1.6 and 5.4 μg C (μg chl a)⁻¹ h⁻¹. Light limitation was found within the mixing depth, while no photoinhibition was observed when surface light was 500 μmol q m⁻² s⁻¹. The mean spectral absorption coefficients of phytoplankton ranged between 0.018 and 0.042 m² (mg chl a)⁻¹ depending on the phytoplankton taxonomy. The quantum yield of photosynthesis varied between 0.027 and 0.076 mol C mol q⁻¹. These high quantum yields are explained by the prevailing high nutrient concentrations in this area. Light intensity plays a major role as limiting factor, even in very shallow water. The phytoplankton close to the surface did not show photoinhibition but had higher UV absorption capabilities.     

Secretion of byssal threads in Mytilus galloprovincialis: quantitative and qualitative values after spawning stress

The effect of spawning events of the mussel Mytilus galloprovincialis on both quantitative and qualitative values of byssus secretion and its associated attachment force was investigated. Byssogenesis rates and absorption efficiency values were significantly reduced after spawning of individuals. However, the maintenance of individuals under sub-optimal conditions (lack of microalgae in the diet) for a week caused no effect on thread’s number. Surprisingly, the attachment force varied within a narrow range of values (1.7–1.9 N) with the exception of a significant drop in the experimental group spawned and kept unfed (1.0 N; P < 0.001), most likely due to a similar pattern of the thread’s thickness variability. Qualitative analysis concerned to the amino acid composition of the byssus highlighted a higher presence of the basic residues histidine and lysine in threads secreted by spawned individuals. The presence of both histidine and lysine residues in the byssal collagen is associated to the formation of cross-links and specifically histidine has a functionality with a pronounced effect on metal chelation to stabilise the integrity of the byssus. Results reported here evidence the necessity to integrate all components that eventually determines the attachment strength of the mussels to get more insight to the plasticity of such secretion. Morphology of the byssus (thickness) secreted under different endogenous conditions of mussels was the major parameter to explain variability in the attachment force. Moreover, aminoacidic composition as quality term of the byssus secreted may also contribute to understand the plasticity of this secretion and needs to be extended in further surveys.     

Response of brown mussel, Perna indica, to elevated temperatures in relation to power plant biofouling control

1. 1. Results of a study on lethal and sublethal responses of different size groups of the tropical brown mussel, Perna indica, when exposed to different temperatures are presented.

2. 2. Exposure to a temperature of 38°C showed 100% mortality of 9 mm size group mussels in 120 min.

3. 3. Mortality was dependent on age (size) of the mussels, young ones being more susceptible than older ones.

4. 4. All size groups showed a progressive reduction in physiological activities such as filtration rate, foot activity and byssus thread production when temperature was increased from 30°C.

5. 5. This study suggests that heat treatment is an attractive alternative to chlorination for mussel fouling control in tropical power stations.

     

Survival and behaviour of juvenile unionid mussels exposed to thermal stress and dewatering in the presence of a sediment temperature gradient

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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.     

Preferences of the Ponto-Caspian amphipod Dikerogammarus haemobaphes for living zebra mussels

A Ponto-Caspian amphipod Dikerogammarus haemobaphes has recently invaded European waters. In the recipient area, it encountered Dreissena polymorpha , a habitat-forming bivalve, co-occurring with the gammarids in their native range. We assumed that interspecific interactions between these two species, which could develop during their long-term co-evolution, may affect the gammarid behaviour in novel areas. We examined the gammarid ability to select a habitat containing living mussels and searched for cues used in that selection. We hypothesized that they may respond to such traits of a living mussel as byssal threads, activity (e.g. valve movements, filtration) and/or shell surface properties. We conducted the pairwise habitat-choice experiments in which we offered various objects to single gammarids in the following combinations: (1) living mussels versus empty shells (the general effect of living Dreissena ); (2) living mussels versus shells with added byssal threads and shells with byssus versus shells without it (the effect of byssus); (3) living mussels versus shells, both coated with nail varnish to neutralize the shell surface (the effect of mussel activity); (4) varnished versus clean living mussels (the effect of shell surface); (5) varnished versus clean stones (the effect of varnish). We checked the gammarid positions in the experimental tanks after 24 h. The gammarids preferred clean living mussels over clean shells, regardless of the presence of byssal threads under the latter. They responded to the shell surface, exhibiting preferences for clean mussels over varnished individuals. They were neither affected by the presence of byssus nor by mussel activity. The ability to detect and actively select zebra mussel habitats may be beneficial for D. haemobaphes and help it establish stable populations in newly invaded areas.     

Byssus attachment strength of two mytilids in mono-specific and mixed-species mussel beds

The mussel Xenostrobus securis is endemic to the brackish waters of New Zealand and Australia, but has successfully invaded the inner Galician Rías of NW Spain, where it coexists with the indigenous mussel Mytilus galloprovincialis. In this laboratory study, the plasticity of the byssus attachment strength of two mytilids was compared by manipulating substratum, salinity, and bed assembly. M. galloprovincialis showed stronger byssus detachment strength than X. securis, despite lower byssus coverage. Both species responded similarly to the substratum, with substantially lower byssus strength on methacrylate, which offered the lowest surface free energy. Byssus detachment values for M. galloprovincialis were lower at lower salinity. In mixed beds, a number of mussels moved upwards, eventually colonising the upper layers of the assemblage. This behaviour increased byssus strength but only for X. securis. X. securis is adapted to a wide spectrum of abiotic conditions, a trait that may promote its dissemination within estuarine environments.