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.     

Patterns of migration between feeding and spawning sites in a coral reef surgeonfish

Many coral reef fishes exhibit regular localised migrations between feeding and spawning areas, but the factors affecting these migration patterns, such as the distance, frequency and spawning site fidelity are poorly understood. The aim of this study was to investigate the patterns of migration to spawning sites of the surgeonfish, Ctenochaetus striatus (Acanthuridae). We explored relationships amongst an individual’s size and sex, the distance and frequency it migrated from its feeding area to spawning sites, fidelity to particular spawning sites and the number of individuals that aggregated to spawn. In order to achieve this, 406 C. striatus were captured and tagged on inshore reefs in Kimbe Bay (5°30′S 150°6′E), New Britain, Papua New Guinea. Tagged individuals were consistently observed within spatially discrete but overlapping feeding areas (maximum diameter averaging <13 m). The mean distance migrated was 58 m (ranging from 2 to 291 m). No tagged individuals were witnessed spawning at more than one site. Whilst most individuals (n = 88) migrated to the spawning site that was closest to their feeding areas, those that migrated to sites further away (n = 9) always spawned at sites where the number of conspecifics aggregating was larger. Neither the size nor the sex of individuals limited migration distance. However, males migrated significantly more frequently than females (on average once every 2 days vs. once every 3 days), and migration frequency was positively correlated with size in females. Migration distance did not affect the frequency with which individuals spawned. Whether patterns of migration are determined by cost-benefit optimisation, tradition, or an alternative mechanism is unknown.     

Clumping behavior and byssus production as strategies for substrate competition in Mytilus edulis

Laboratory experiments showed that the mussel Mytilus edulis aggregated more intensely around living organisms (the bivalve Hiatella arctica and the solitary ascidian Styela rustica, which commonly co‐occur with mussels in fouling communities) than around inanimate objects. When exposed to an inanimate object, mussels attached their byssal threads primarily to the substrate, close to the object, but when exposed to a living organism, they attached their byssal threads directly to the organism. The ascidian was more intensely covered with byssal threads than was the bivalve. Mussel attachment to the ascidians was apparently determined by the physical characteristics of the tunic and to a lesser extent by the excretion‐secretion products released by S. rustica. This study indicates that mussels can use byssus threads as a means of entrapment of potential competitors for space. It remains unclear why mussels preferentially attached to ascidians compared to the bivalve. This can be explained either by competitive interactions, or by attractiveness of the ascidian tunic as an attachment substratum.     

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.     

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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Green mussel Perna viridis L.: attachment behaviour and preparation of antifouling surfaces

The green mussel Perna viridis LINNE can be kept in simulated seawater for more than 6 months in good condition. The mussel forms many threads by secreting an adhesive protein from the foot, and attaches with more than 50 byssal threads, which makes most mussels clump together. In order to investigate the preparation of the antifouling surfaces toward green mussels, the attachment of mussels was tested using glass surfaces modified with silane coupling agents, together with non-treated material surfaces such as glass and silicone. The correlation between the attachment percentage and the mean number of the secreted byssus was highly significant, indicating that the mussel selects a favorable surface prior to the secretion of byssus. The relationships between the mussel attachment and the surface chemical parameters (surface free energy (sfe) and its dispersion and polar components) were examined based on a working hypothesis, which we have previously reported. The result of statistical regression test indicated that a certain correlation was found between the dispersion component and the mussel attachment, while the polar component did not correlate to the mussel attachment. The present surface chemical approach provided an additional clue for the preparation of ecologically clean antifouling materials that takes into account the combination of the wettability of both the marine adhesive proteins (MAP) and the modified surfaces.     

Activation of spawning in zebra mussels by algae-, cryptomonad-, and gamete-associated factors

In zebra mussels (Dreissena polymorpha) simultaneous release of gametes and peaks in larval densities at particular locations suggest that spawning is triggered by synchronizing stimuli. Furthermore, spawning tends to occur only after an adequate environmental temperature is reached. To test the hypothesis that phytoplankton and gamete-associated chemicals initiate spawning in zebra mussels and that the responsiveness to such chemicals is affected by ambient temperature, the spawning response of zebra mussels to extracts from algae, a cryptomonad, and a cyanobacterium and to water associated with released gametes was assayed in animals acclimated to 12 ^C and 17 ^C. For animals held at 12 ^C, only serotonin, a known activator of bivalve spawning used as a positive control, stimulated spawning. However, for animals acclimated to 17 ^C, extracts made from a diatom (Phaeodactylum), a brown alga (Fucus), and a cryptomonad (Rhodomonas) stimulated spawning in both sexes; extracts from green algae (Platymonas and Dunaliella) and a cyanobacterium (Oscillatoria) did not cause spawning. Water associated with either released sperm or eggs elicited spawning in both females and males. Positive controls, stimulated with serotonin, spawned at a high (>90%) rate, whereas no negative control spawned. Thus, phytoplankton chemicals and gamete-associated factors may have a role in synchronizing spawning in zebra mussels once adequate ambient temperature is reached.     

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。     

Reproductive cycle and reproductive output of the sea urchin Loxechinus albus (Echinodermata: Echinoidea) from Beagle Channel, Tierra del Fuego, Argentina

Reproductive cycle, frequency and duration of spawning, energetic content of gonads, and reproductive output of the common green sea urchin Loxechinus albus were analyzed in the Beagle Channel (Tierra del Fuego) between May 2004 and May 2005. Gonad indices (GI, percentages of gonad mass in total body mass) were significantly higher in March, April, July, and August than in November and May, thus showing a negative correlation with the photoperiod. Highest GI values of mature individuals were observed in August, and spawning occurred from September to December. In females, the mass-specific energy content of gonads (ECG) was highest in spawned gonads and lowest in mature ones, while in males ECG values were higher in immature stage and lower in premature and mature stages. High ECG values can be explained by the abundance of nutritive phagocytes. Both ECG and total gonad energy content (TECG) were higher in females than in males. Mean reproductive output was 7.28% for females and 6.15% for males (expressed as the difference between mean GI of mature and spawned gonads) and 25.02 kJ for females and 19.26 kJ for males (expressed as the difference between mean TECG of mature and spawned gonads).     

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)     

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.     

Interspecific variation in spawning time and male mating tactics of the parrotfishes on a fringing coral reef at Iriomote Island,Okinawa

Spawning time and male mating tactics of parrotfishes (family Scaridae) were investigated on a fringing coral reef at Iriomote Island, Okinawa. Spawning was observed in 14 species, and more frequently in more abundant species such as Chlorurus sordidus, Scarus rivulatus and Chlorurus bowersi. At the reef-edge spawning site, C. bowersi spawned at high tide, C. sordidus spawned both at high tide and in the early morning, whereas Calotomus carolinus and most of the Scarus species such as S. rivulatus spawned only in the early morning, mostly 0630–0830 h. Spawning only in the early morning irrespective of tide phase and moon age has seldom been reported from the scarid species of other localities. It is suggested that spawning in the early morning would be adaptive in species such as S. rivulatus, which migrated considerable distances (ca. 500 m) to the inshore feeding sites, in order to minimize feeding losses due to migration. For male mating tactics, pair spawning by territorial TP (terminal phase) males occurred in all 14 species, and streaking and group spawning by nonterritorial small IP (initial phase) males were seen more frequently in more abundant species. Moreover, group spawning by nonterritorial TP males, which were larger than the IP males but smaller than the territorial TP males, frequently occurred in S. rivulatus. Such mating tactics of TP males have not been reported from Scaridae.     

Zebra mussel adhesion: structure of the byssal adhesive apparatus in the freshwater mussel, Dreissena polymorpha

The freshwater zebra mussel (Dreissena polymorpha) owes a large part of its success as an invasive species to its ability to attach to a wide variety of substrates. As in marine mussels, this attachment is achieved by a proteinaceous byssus, a series of threads joined at a stem that connect the mussel to adhesive plaques secreted onto the substrate. Although the zebra mussel byssus is superficially similar to marine mussels, significant structural and compositional differences suggest that further investigation of the adhesion mechanisms in this freshwater species is warranted. Here we present an ultrastructural examination of the zebra mussel byssus, with emphasis on interfaces that are critical to its adhesive function. By examining the attached plaques, we show that adhesion is mediated by a uniform electron dense layer on the underside of the plaque. This layer is only 10-20 nm thick and makes direct and continuous contact with the substrate. The plaque itself is fibrous, and curiously can exhibit either a dense or porous morphology. In zebra mussels, a graded interface between the animal and the substrate mussels is achieved by interdigitation of uniform threads with the stem, in contrast to marine mussels, where the threads themselves are non-uniform. Our observations of several novel aspects of zebra mussel byssal ultrastructure may have important implications not only for preventing biofouling by the zebra mussel, but for the development of new bioadhesives as well.     

The timing of spawning in capelin (Mallotus villosus Müller) at a coastal location in eastern Newfoundland

We tested three hypotheses concerning the timing of spawning for a circumpolar species, capelin (Mallotus villosus), for which timing of larval emergence is known to be synchronized by physical conditions. The first hypothesis, developed from previous studies, was that spawning would be synchronized by upwelling events. Initial results from Middle Cove Beach in eastern Newfoundland indicated that spawning was not synchronized with upwelling. We next hypothesized that spawning was a function of several environmental variables. Results from logistic regression indicated that neither single-factor nor multi-factor models could explain the timing of spawning. Single variables could predict spawning in some years but no variable could reliably predict the time of spawning year after year. Finally, we hypothesized that the probability of spawning increased as a set of significant variables approached preferred levels. For capelin at Middle Cove, the set of variables that influence capelin spawning were identified as wave height, sea surface roughness and capelin abundance in the water. Thus only a combination of variables explained the timing of spawning for capelin. Preferred conditions for capelin spawning were wave heights less than 20 cm at the beach, a sea surface with a slight ripple, and an intermediate rank abundance of capelin in the water corresponding to hundreds to thousands of individuals. Capelin abundance alone was not a useful predictor. During the course of the study we observed a shift in the dates that capelin arrived and spawned at the beach. During 1987–1990 capelin spawned at Middle Cove Beach during June, but in more recent years (1991–1993) capelin did not arrive or spawn until July.     

Algal-induced spawning in the marine mussel Mytilus californianus 4

The characteristics of algal-induced spawning of the marine mussel Mytilus californianus were studied. Exposure of mature individuals to culture suspensions of the unicellular alga Pseudoisochrysis paradoxa elicited copious and synchronous release of gametes. Alkaline conditions were necessary to make the animals responsive to the spawning stimulus provided by the algae. The filtered, cell-free fraction of the algal suspensions also stimulated spawning, suggesting that an active principle is secreted into the culture media by the algae.

A key requirement for the development of an efficient and economically viable molluscan mariculture program is the capability to induce copious, synchronous spawning of gravid individuals. This procedure should be easily managed and have no deleterious effects on the gametes, fertilization, or the ontological development of the animals. Miyazaki [1] induced spawning in male oysters by exposing them to an extract of green algae. Recently it was reported that the marine mussels, M. californianus, could be induced to spawn by exposing them to cultures of the marine algae, P. paradoxa [2], This paper summarizes our efforts to characterize this spawning response of mussels to high concentrations of cultured marine algae.     

Morphology of the Bivalve Mollusks Crenomytilus grayanus and Mytilus coruscus in Relation to Their Spatial Distribution in the Upper Subtidal Zone

The morphology of the shell and byssus threads was studied in two closely related mussel species Crenomytilus grayanus and Mytilus coruscus. The two species differ significantly from each other in the shell shape and in the degrees of development and deformation of byssus threads. These differences, in turn, determine (either directly or indirectly) the differences in strength of the byssal attachment and are discussed in terms of their functional morphology with respect to the spatial distribution of the mussels in marine coastal zones.     

Plasticity of fertilization rates under varying temperature in the broadcast spawning mussel,Mytilus galloprovincialis

Oceans are a huge sink for the increased heat associated with anthropogenic climate change, and it is vital to understand the heat tolerance of marine organisms at all life stages to accurately predict species’ responses. In broadcast spawning marine invertebrates, reproduction is a vulnerable process in which sperm and eggs are released directly into the open water. Gametes are then exposed to fluctuating environmental conditions that may impact their fertilizing capacity. Using the broadcast spawning Mediterranean mussel, Mytilus galloprovincialis, as a model species, we performed blocks of factorial mating crosses to assess the variance in fertilization rates among individuals under both ambient and elevated temperatures. Overall, we found a small, but significant decline in fertilization rates with elevated temperatures. However, there was substantial plasticity in responses, with particular mussels having increased fertilization under elevated temperatures, although the majority showed decreased fertilization rates. Our results suggest possible future reproductive costs to ocean warming in M. galloprovincialis, although it is also possible that genetic variation for thermal sensitivity may allow for adaptation to changing environmental conditions.     

Observations of spawning in the Leather Bass, <Emphasis Type="Italic">Dermatolepis dermatolepis</Emphasis> (Teleostei: Epinephelidae), at Cocos Island,Costa Rica

Courtship and spawning behavior is described for the Leather Bass, Dermatolepis dermatolepis, from observations made from a manned submersible at Cocos Island, Costa Rica on 19 days between November 2006 and February 2007. Spawning occurred in the evening (16:00–18:30 h) in subgroups of 10–32 individuals that formed within aggregations of 50–70 individuals. The mating sequence was characterized by subgroup formation and crowding behavior followed by a brief vertical spawning rush, gamete release, and the rapid separation of participants. The aggregation was present on all dives, and spawning was not restricted to specific lunar phases.     

Migration of green sturgeon, Acipenser medirostris, in the Sacramento River

Adult green sturgeon, Acipenser medirostris, were collected in San Pablo Bay, California, and surgically implanted with ultrasonic acoustic tags from 2004 to 2006. An array of automated acoustic monitors was maintained in the Sacramento River to record movements of these fish. We presumed movements to known spawning areas (based on previous green sturgeon egg collections) or areas with potential spawning habitat (characterized by substrate, flow, and temperature criteria) represented a “spawning migration.” Three separate annual “spawning migrations” were recorded involving 15 individuals. The majority of the Sacramento River migrants entered the system in the months of March and April. Two different patterns of “spawning migration” and out-migration were observed. Six individuals potentially spawned, over-summered and moved out of the river with the first fall flow event. This is believed to be the common behavior of the green sturgeon. Alternatively, nine individuals promptly moved out of the Sacramento River before 1 September, and any known flow or temperature cue. Some green sturgeon appeared to be impeded on their upstream movement by the 15 May closure of the Red Bluff Diversion Dam, and at least five passed under the dam gates during downstream migration. A delay in the closure of the Red Bluff Diversion Dam would likely allow upstream passage of spawning green sturgeon, further, the potential mortality affects of downstream passage beneath the Red Bluff Diversion Dam should be assessed. Specific protection should be also given to the large aggregation of green sturgeon located in the reach of the Sacramento River adjacent to the Glen Colusa Irrigation District pumping facility.