Predator and prey: the role of the round goby Neogobius melanostomus in the western Baltic

Different studies on the position of the non-indigenous species Neogobius melanostomus within the coastal food web of the Pomeranian Bay (western Baltic) were performed, resulting in a quantitative and qualitative species list of prey organisms found in the stomachs of the invader and an estimation concerning the importance of round goby as prey for different resident predators. It seems that the colonization process is not fully completed yet, but the results reveal that the species is already established in the food web 16 years after the first observation within the study area. The results show that N. melanostomus feed upon a wide range of different resident organisms. While a direct predation effect on native fish species appears rather unlikely, indirect effects such as competition cannot yet be excluded. In addition, our results reveal an ontogenetic diet shift and that the round goby itself already serves as an important prey for piscivorous fish and seabirds. Finally, we formulate different hypotheses based on our results which will require further research.     

Expansion and fragment settlement of the non-native seagrass Halophila stipulacea in a Caribbean bay

The non-native seagrass species Halophila stipulacea has spread throughout the Eastern Caribbean since 2002, and could potentially impact the functioning of local seagrass ecosystems. Important characteristics for invasiveness, such as dispersal, recruitment and expansion of H. stipulacea at a local scale, are unknown. We assessed H. stipulacea expansion rates within Lac Bay, Bonaire, Dutch Caribbean (7?km²), since its establishment in 2010 and tested the settlement potential of uprooted vegetative fragments of H. stipulacea. Using 49 fixed locations, we observed that between 2011 and 2015 the occurrence of H. stipulacea in the bay increased significantly from 6% to 20% while native Thalassia testudinum occurrence decreased significantly from 53% to 33%. Free-floating H. stipulacea fragments that were collected and tethered above the sediment rooted within 10 days with a settlement success rate of 100%. The growth of settled fragments was on average 0.91 shoots d^?1. The ongoing shift from native T. testudinum to introduced H. stipulacea dominated meadows may have important consequences for multiple Caribbean seagrass ecosystem functions. Given the large difference in size between the two seagrass species, functions such as coastal protection, habitat structure, food availability, and the stability and resilience of these systems can be altered. The next steps towards modelling future expansion of H. stipulacea throughout the Caribbean and beyond should include the assessment of fragment viability and dispersal distance, and the impacts of natural and anthropogenic disturbance on vegetative fragment density, dispersion and settlement by this species.     

Heat-shock response of the upper intertidal barnacle Balanus glandula: thermal stress and acclimation

In the intertidal zone in the Pacific Northwest, body temperatures of sessile marine organisms can reach 35 degrees C for an extended time during low tide, resulting in potential physiological stress. We used immunochemical assays to examine the effects of thermal stress on endogenous Hsp70 levels in the intertidal barnacle Balanus glandula. After thermal stress, endogenous Hsp70 levels did not increase above control levels in B. glandula exposed to 20 and 28 degrees C. In a separate experiment, endogenous Hsp70 levels were higher than control levels when B. glandula was exposed to 34 degrees C for 8.5 h. Although an induced heat-shock response was observed, levels of conjugated ubiquitin failed to indicate irreversible protein damage at temperatures up to 34 degrees C. With metabolic labeling, we examined temperature acclimation and thermally induced heat-shock proteins in B. glandula. An induced heat-shock response of proteins in the 70-kDa region (Hsp70) occurred in B. glandula above 23 degrees C. This heat-shock response was similar in molting and non-molting barnacles. Acclimation of B. glandula to relatively higher temperatures resulted in higher levels of protein synthesis in the 70-kDa region and lack of an upward shift in the induction temperature for heat-shock proteins. Our results suggest that B. glandula may be well adapted to life in the high intertidal zone but may lack the plasticity to acclimate to higher temperatures.     

Early phase of the invasion of Balanus glandula along the coast of Eastern Hokkaido: changes in abundance,distribution, and recruitment

To understand the patterns and processes associated with the population dynamics of Balanus glandula during the early phase of invasion along the Pacific coast of eastern Hokkaido, population surveys were conducted from 2002 to 2011 at five shores, each consisting of five paired plots (scraped recruitment plot and unscraped establishment plot), along 49 km of coastline located 144 km east of the eastern front of the invasion of this species in 2000. Larval recruitment was first detected in 2004, but the establishment of a population was not observed until 2 years later at the westernmost shore of the study area. Occurrence increased from non-native barnacle present in 4 % of plots in 2006 to 100 % in 2011, but mean coverage remained low (<5 %) in 2011. Most local population coverage fluctuated without indicating clear temporal trends, but coverage in one plot showed a consistent pattern of rapid increase. Local extinctions occurred, but rates of local extinction decreased with time as larval recruitment increased. Lag times between recruitment and establishment occurred for 64 % of the paired plots and ranged from 1 to 4 years. Lag times decreased after 5 years, when larval recruitment increased. These findings suggest that the intensity of larval recruitment determined invasion dynamics during this early phase of the invasion, and the monitoring of recruitment is therefore essential for early detection of invasions by sessile marine organisms and prediction of their range expansion.     

Analysis of behavioural rejection of micro-textured surfaces and implications for recruitment by the barnacle Balanus improvisus

Experiments performed in the field and in the laboratory show that the barnacle, Balanus improvisus, preferentially settles on smooth surfaces. Settlement and recruitment of B. improvisus was evaluated on micro-textured surfaces with scales of surface texture ranging from 1 to 100 μm in profile heights. Surface texture with profile heights within a topographic range of 30-45 μm reduced settlement and recruitment by 92% as compared to smooth surfaces. The reduction in recruitment on micro-textured surfaces is best explained by behavioural responses to surface topography. Behavioural experiments show that cyprids have a higher propensity for smooth surfaces than for micro-textured surfaces. Cyprids spend more time exploring smooth surfaces and more time swimming when exposed to micro-textured surfaces. Micro-textured surfaces are more often rejected by cyprids after exploration than smooth surfaces. It is suggested that some scales of surface texture could be exploited to improve future anti-fouling techniques in geographical areas where Balanus improvisus is a severe fouling problem.     

Linking stages of life history: How larval quality translates into juvenile performance for an intertidal barnacle (Balanus glandula)

Many marine invertebrates with complex life cycles produce planktoniclarvae that experience environmental conditions different fromthose encountered by adults. Factors such as temperature andfood, known to impact the larval period, can also affect larvalsize and consequently the size of newly settled juveniles. Afterdocumenting natural variation in the size of cyprids (the finallarval stage) of the barnacle Balanus glandula, we experimentallymanipulated temperature and food given to larvae to producecyprids of differing sizes but within the size range of cypridsfound in the field. In a set of trials in which larvae of B.glandula were raised on full or reduced rations in the laboratoryand subsequently outplanted into the field as newly metamorphosedjuveniles, we explored the effects of larval nutrition and sizeon juvenile performance. Larvae that received full rations throughouttheir feeding period produced larger cyprids (with more lipidand protein). These larger cyprids grew faster as juvenilesand sometimes survived better in the field than juveniles fromlarvae that had their food ration reduced in the last feedinginstar. For naturally settling barnacles brought into the laboratorywithin 2 days of settlement and fed, we found that initial juvenilesize was a good predictor of juvenile size even after 2 weeksof growth. By manipulating food given to juveniles that werederived from larvae fed either full or reduced rations, we foundthat larval nutritional effects persisted in juveniles for 2–3times the period that larvae experienced altered food rations.     

Modeling the effects of environmental variability on Balanus glandula larval development

Planktonic larvae encounter variability in the marine environmentas they develop, and rates of larval development are affectedby changes in certain environmental variables. We examine theeffects of variation in environmental parameters on the durationof the naupliar period of Balanus glandula larvae. Naupliardevelopment rate is modeled as a function of temperature andfood concentration. Simulations of the time course of developmentwere driven by time series of temperature and food concentrationcollected at two locations on the US west coast. Predicted naupliardurations are reasonable in comparison to previous observations.Simulations of larval development and dispersal were drivenby output of a three-dimensional physical circulation modeland remotely sensed chlorophyll. These simulations demonstratethe impact of spatial variation in current velocity, temperatureand chlorophyll on the duration of the larval period, the dispersaltrajectory of these organisms and ultimately on recruitmentsuccess. Our naupliar development model coupled with a physicalcirculation model is a useful tool for addressing questionsof intertidal population ecology.     

The lateral photoreceptor of the barnacle,Balanus eburneus

Summary The lateral eye of the barnacle, Balanus eburneus, fixed in highly concentrated osmium is a lens-shaped body of approximately 250 m in diameter and about 75 m thick. It contains three photoreceptor cells which occupy about 42% of its volume. The photoreceptor cells are irregularly shaped and extend countless dendritic processes which bear rhabdomeres at their ends. Individual rhabdomeres come into contact with rhabdomeres originating from dendrites of the same or of one of the other visual cells. Thirteen per cent of the volume of the photoreceptor cells is taken up by the rhabdomeres. The membranes of the rhabdomeric microvilli contain globular subunits which suggest a 70 Å spacing of rhodopsin molecules. There are two kinds of glial cells. One kind, type A glial cells, makes contact with the fibrous capsule of the photoreceptor. The other kind, type B glial cells, is associated with the photoreceptor cells and extends countless tiny cytoplasmic extensions which interdigitate with similar extensions of the receptor cells. There are approximately 95 type B glial cells and 130 type A glial cells in the receptor. The cytoplasm of the photoreceptor cells contains countless small Golgi fields, mitochondria, microtubules, multivesicular and multilamellar bodies. The extracellular space of the photoreceptor is less than 0.1% of its total volume.The authors wish to thank Mrs. G. Theisen and Miss D. Hupp for expert technical assistance and Drs. M. Behrens, C. Helrich, and C.C. Krischer for many inspiring discussions. This study was partly supported by the SFB 160 of the Deutsche Forschungsgemeinschaft     

Variation among families for characteristics of the adhesive plaque in the barnacle Balanus amphitrite

A quantitative genetics approach was used to examine variation in the characteristics of the adhesive plaque of the barnacle Balanus amphitrite Darwin attached to two silicone substrata. Barnacles settled on silicone polymer films occasionally form thick, soft adhesive plaques, in contrast to the thin, hard plaques characteristic of attachment to other surfaces. The proportion of barnacles producing a thick adhesive plaque was 0.31 for Veridian, a commercially available silicone fouling-release coating, and 0.18 for Silastic T-2, a silicone rubber used for mold-making. For both materials, significant variation among maternal families in the proportion of barnacles producing a thick adhesive plaque was observed, which suggests the presence of genetic variation, or maternal environmental effects, for this plaque characteristic. For the Veridian coating, barnacles expressing the thick adhesive plaque also exhibited significantly reduced tenacity. This represents the first reported case for potential genetic control of intraspecific phenotypic variation in the physical characteristics and tenacity of the adhesive of a fouling invertebrate.     

Evidence for different pharmacological targets for imidazoline compounds inhibiting settlement of the barnacle Balanus improvisus

We describe the effect of eight different imidazoline/guanidinium compounds on the settlement and metamorphosis of larvae of the barnacle Balanus improvisus. These agents were chosen on the basis of their similar pharmacological classification in vertebrates and their chemical similarity to medetomidine and clonidine, previously described as highly potent settlement inhibitors (nanomolar range). Seven of the tested compounds were found to inhibit settlement in a dose-dependent manner in concentrations ranging from 100 nM to 10 microM without any significant lethal effects. In vertebrate systems these substances have overlapping functions and interact with both alpha-adrenoceptors as well as imidazoline binding sites. Antagonizing experiments using the highly specific alpha(2)-antagonist methoxy-idazoxan or agmatine (the putative endogenous ligand at imidazoline receptors) were performed to discriminate between putative pharmacological mechanisms involved in the inhibition of cyprid settlement. Agmatine was not able to reverse the effect of any of the tested compounds. However, methoxy-idazoxan almost completely abolished the settlement inhibition mediated by guanabenz (alpha(2)-agonist, I(2) ligand), moxonidine (alpha(2)-agonist, I(1) ligand) and tetrahydrozoline (alpha-agonist, I(2) ligand). The actions of cirazoline (alpha(1)-agonist, I(2) ligand) BU 224 (I(2) ligand) and metrazoline (I(2) ligand) were not reversed by treatment with methoxy-idazoxan. These results suggest that the settlement inhibition evoked by the I(2) ligands and alpha(2)-agonists used in this study of the neurologically simple but well-organized barnacle larva is mediated through different physiological targets important in the overall settlement process.     

Spatial pattern in a low-density population of the barnacle Balanus amphitrite Darwin

The spatial distribution of an intertidal population of the barnacleBalanus amphitrite was studied close to a sewage outfall nearQuequén, Argentina (38° 34S,58° 38 30 W). All individuals within an areaof 0.8 m² were mapped during 12 censuses, from September 1990to May 1993. Population density within the study area varied between 401 and99 ind m^-2. Spatial pattern was analysed using mean distancesto the nearest neighbour (NND), goodness of fit tests between observed andexpected frequency distributions of NND, and analysis of crowding. Thespatial pattern of the population was clumped during most of the studyperiod. This trend persisted, although not reaching statisticalsignificance, when the population density decreased as a consequence ofmortality and failure of successive annual recruitments. The cohortrecruited in summer 1992 (49 individuals) was randomly distributed relativeto adult barnacles already present within the study area. No consistentrelationship was observed between mortality and NND.     

Base plate mechanics of the barnacle Balanus amphitrite (=Amphibalanus amphitrite)

The mechanical properties of barnacle base plates were measured using a punch test apparatus, with the purpose of examining the effect that the base plate flexural rigidity may have on adhesion mechanics. Base plate compliance was measured for 43 Balanus amphitrite (=Amphibalanus amphitrite) barnacles. Compliance measurements were used to determine flexural rigidity (assuming a fixed-edge circular plate approximation) and composite modulus of the base plates. The barnacles were categorized by age and cement type (hard or gummy) for statistical analyses. Barnacles that were 'hard' (> or =70% of the base plate thin, rigid cement) and 'gummy' (>30% of the base plate covered in compliant, tacky cement) showed statistically different composite moduli but did not show a difference in base plate flexural rigidity. The average flexural rigidity for all barnacles was 0.0020 Nm (SEM +/- 0.0003). Flexural rigidity and composite modulus did not differ significantly between 3-month and 14-month-old barnacles. The relatively low flexural rigidity measured for barnacles suggests that a rigid punch approximation is not sufficient to account for the contributions to adhesion mechanics due to flexing of real barnacles during release.     

Roles of dopamine and serotonin in larval attachment of the barnacle, Balanus amphitrite

In order to clarify the roles of neurotransmitters including serotonin and dopamine in larval settlement (attachment) and metamorphosis of the barnacle Balanus amphitrite, the effects of lisuride, which acts as both a serotonin agonist/antagonist and a dopamine agonist, were examined. Lisuride did not induce larval attachment and metamorphosis; however, it promoted only larval behavior of searching for attachment sites without actual attachment to substrata which lasted for 5 to 6 days in a dose-dependent manner. Further evidence was obtained with a range of agonists/antagonists; serotonin agonists promoted the attachment, while serotonin antagonists inhibited it. Similarly, dopamine agonists inhibited the attachment. Furthermore, mixtures of serotonin and dopamine showed similar effects to those of lisuride. These results suggested that the promotion effect on larval searching behavior was derived from a combination of activities of serotonin and dopamine. Moreover, both serotonin and dopamine were detected in cyprids by HPLC. Thus, larval attachment process is regulated by both serotonin and dopamine neurons in this species. J. Exp. Zool. 284:746-758, 1999. Copyright 1999 Wiley-Liss, Inc.     

Eicosanoid generation in the intertidal barnacle, Balanus perforatus--effect of season and reproductive status

The nature and quantity of the principal lipoxygenase (LO) products generated by the intertidal barnacle Balanus perforatus were examined at monthly intervals and their potential involvement in reproductive events was investigated. The main mono-hydroxy products generated were found to be formed through the action of an 8-lipoxygenase (8-LO) activity and were the mono-hydroxy fatty acids, 8-hydroxyeicosapentaenoic acid (HEPE) and 8-hydroxyeicosatetraenoic acid. Generation of these products was found to be correlated with the environmental seawater temperature, although no change in either 8-LO activity or the precursor fatty acid levels in total phospholipids was found with the time of the year. Changes in fatty acid composition measured in animals collected from summer and winter conditions were found to follow the pattern expected by homeoviscous adaptation of a cold-acclimated animal. Oogenesis was found to occur in August and was linked to a significant reduction in 8-HEPE generation. Spermatozoa were found to be present year-round in the seminal vesicles although the testes became atrophic during the winter months.     

Characterisation of the bacteria associated with barnacle,Balanus amphitrite,shell and their role in gregarious settlement of cypris larvae

The acorn barnacle Balanus amphitrite (syn. Amphibalanus amphitrite) is a model organism to investigate pelago-benthic transitions in marine invertebrates. A driver for larval settlement in this organism is the need to attach close to conspecifics, to allow reproduction to take place. Adult barnacles are covered by microbial biofilms and the contribution of these biofilms to conspecific recognition is not fully understood. Little information is available on microbial communities associated with B. amphitrite. We compared biofilm communities from the barnacle shell surface with those from the surrounding rocks using the culture-independent methods of quantitative PCR and denaturing gradient gel electrophoresis. Quantification of the relative abundances of higher bacterial taxa showed that barnacles hosted a greater proportion of α-Proteobacteria compared to rock-associated biofilms (p < 0.01). Differences in relative abundances of other taxa were not observed but DGGE profiling suggested that differences were present at lower taxonomic levels. The capacity of these communities to influence larval settlement was assessed by growing multispecies biofilms on artificial medium, obtained by extracting nutrients from adult barnacles. Biofilms composed of shell-associated bacteria were capable of promoting conspecific settlement by 67% compared to control surfaces (p < 0.05), while rock-associated communities showed contrasting effects. A taxonomic comparison of settlement-stimulating and -inhibiting bacteria was performed by DGGE and band sequencing. All partial 16S rRNA genes sequenced were similar to members of the Vibrio and Pseudoalteromonas genera, suggesting that larvae can detect and respond to variations in the composition of microbial biofilms at low taxonomic levels. Our results indicate that barnacle larvae may be able to detect parentally-associated biofilms and use this information to settle close to members of its own species.     

Antifouling activity of simple synthetic isocyanides against larvae of the barnacle Balanus amphitrite

Twelve simple linear isocyanides were synthesized and examined for antifouling activity and toxicity against cyprid larvae of the barnacle, Balanus amphitrite. Larval settlement was inhibited, with EC50 values of 0.046-1.90 microg ml(-1), and they were much less toxic (LD50 values ranging over 21.28 microg ml(-1)) than CuSO4 (EC50 0.30 microg ml(-1) and LD50 2.95 microg ml(-1)). The data indicate that simple linear isocyanides are promising non-toxic antifouling agents.     

The complex barnacle perfume: identification of waterborne pheromone homologues in Balanus improvisus and their differential expression during settlement

A key question in barnacle biology is the nature of cues that induce gregarious settlement. One of the characterised cues is the waterborne settlement pheromone (WSP). This study aimed to identify WSP homologues in Balanus improvisus and to investigate their expression during settlement. Six WSP homologues were identified, all containing an N-terminal signal peptide, a conserved core region, and a variable C-terminus comprising several -GR- and -HDDH- motifs. The B. improvisus WSP homologues were expressed in all settlement stages but showed different expression patterns. The homologue most similar to the B. amphitrite WSP was the most abundant and was constantly expressed during settlement. In contrast, several of the other WSP homologues showed the greatest expression in the juvenile stage. The presence of several WSP homologues suggests the existence of a pheromone mix, where con-specificity might be determined by a combination of sequence characteristics and the concentration of the individual components.