Host location in flow by larvae of the symbiotic barnacle Trevathana dentata using odour-gated rheotaxis

The detection and location of specific organisms in the aquatic environment, whether they are mates, prey or settlement sites, are two of the most important challenges facing aquatic animals. Large marine invertebrates such as lobsters have been found to locate specific organisms by navigating in the plume of chemicals emitted by the target. However, active plume tracking in flow by small organisms such as marine larvae has received little scientific attention. Here, we present results from a study examining host location in flow by nauplius larvae of the barnacle Trevathana dentata, which inhabits the stony reef coral Cyphastrea chalcidicum. The experiments included analysis of larval motion in an annular flume under four conditions: (i) still water, (ii) in flow, (iii) in still water with waterborne host metabolites and (iv) in flow with host metabolites. Our results show that T. dentata nauplii are unable to locate their target organism in still water using chemotaxis, but are capable of efficient host location in flow using odour-gated rheotaxis. This technique may enable host location by earlier, less-developed larval stages.     

Hydrodynamics of larval settlement from a larva's point of view

Many benthic marine invertebrate animals release larvae that are dispersed by ocean currents. These larvae swim and can respond to environmental factors such as chemical cues. However, larvae are so small (generally 0.01-1 mm) that they are often assumed to be passive particles whose trajectories are determined by the motion of the water in which they are riding. Therefore, marine larvae are useful model organisms to study the more general question of how the locomotion of very small animals in complex, variable natural habitats is affected by the motion of the fluid (water or air) around them. Studying larval locomotion under conditions of water flow encountered in nature is challenging because measuring the behavior of an individual microscopic organism requires high magnification imaging that is difficult to do in the field. The purpose of this article is to synthesize in one place the various approaches that we have been using to address the technical challenges of studying the locomotion of microscopic larvae in realistic ambient flow. The steps in our process include: (1) measuring water flow in the field; (2) mimicking realistic water movement in laboratory flumes to measure larval scale fluctuations in velocity of flow and concentration of chemical cues; (3) mimicking fine scale temporal patterns of larval encounters with a dissolved chemical cue to record larval responses; (4) using individual-based models to put larvae back into the larger scale environmental flow to determine trajectories; and (5) mimicking fine scale spatial and temporal patterns of larval encounters with water velocities and shear to determine the instantaneous forces on larvae. We illustrate these techniques using examples from our ongoing research on the settlement of larvae onto fouling communities and from our published work on settlement of larvae onto coral reefs. These examples show that water velocities and concentrations of chemical cues encountered by microscopic organisms can fluctuate in fractions of a second and vary over scales of less than a millimeter.     

The importance of flow and settlement cues to larvae of the abalone, Haliotis rufescens Swainson

In flow tank experiments, I tested the relative importance of active and passive processes to larvae settling on manufactured casts that were hydrodynamically rough at a small scale (mm to <1 cm). I predefined two distinct regions of small-scale flow that I used to manipulate larval settlement behaviour of the red abalone Haliotis rufescens Swainson. The larvae show a stringent settlement response associated with coralline red algae. Haliotis rufescens larvae settled preferentially to an inducer regardless of the flow conditions, as expected. However, the ability of H. rufescens larvae to show this stringent behaviour was altered by changing the small-scale flow. When the free-stream velocity was low, the larvae responded to a settlement cue regardless of the small-scale hydrodynamics. When free-stream velocity was higher, the larvae acted increasingly as passive particles in their deposition, but settled only in response to an inducer. The results were consistent in two flow tanks, across 2 years and between different batches of larvae.     

Mini review: hydrodynamics of larval settlement into fouling communities

Flowing water delivers planktonic larvae to surfaces, but also dislodges them. This paper reviews experiments in the field and in laboratory flumes, as well as mathematical models, which have revealed how the interaction of ambient water motion with a developing fouling community affects larval settlement. Although mean current velocities across fouling communities in harbours are low, instantaneous velocities can be much higher due to turbulence and to the velocity oscillations of wind chop and ship wakes. As a fouling community develops, its topography becomes more complex and the range of flow microhabitats on the spatial scale of larvae increases. In spite of the prevalence of waves in shallow coastal habitats, and in spite of the importance to settlement of the fine-scale instantaneous velocities encountered by larvae, most studies of flow effects on larval settlement have focused on unidirectional currents and on temporally- and spatially-averaged aspects of the flow.     

Mini review: Hydrodynamics of larval settlement into fouling communities

Abstract

Flowing water delivers planktonic larvae to surfaces, but also dislodges them. This paper reviews experiments in the field and in laboratory flumes, as well as mathematical models, which have revealed how the interaction of ambient water motion with a developing fouling community affects larval settlement. Although mean current velocities across fouling communities in harbours are low, instantaneous velocities can be much higher due to turbulence and to the velocity oscillations of wind chop and ship wakes. As a fouling community develops, its topography becomes more complex and the range of flow microhabitats on the spatial scale of larvae increases. In spite of the prevalence of waves in shallow coastal habitats, and in spite of the importance to settlement of the fine-scale instantaneous velocities encountered by larvae, most studies of flow effects on larval settlement have focused on unidirectional currents and on temporally- and spatially-averaged aspects of the flow.     

Rapid behavioral responses of an invertebrate larva to dissolved settlement cue

Larvae of the nudibranch Phestilla sibogae were used to study whether a natural dissolved settlement cue (from their prey, Porites compressa, an abundant coral on Hawaiian reefs) induces behavioral responses that can affect larval transport to suitable settlement sites. As cue and larvae are mixed in the turbulent flow over a reef, cue is distributed in fine-scale filaments that the larva experiences as rapid (seconds) on/off encounters. To examine larval responses in this setting, individual larvae were tethered in a small flume with flow simulating water velocity relative to a freely swimming larva, and their responses to realistic temporal patterns of cue encounter were videotaped. Competent larvae quickly ceased swimming in cue filaments and resumed swimming after exiting filaments. The threshold cue concentration eliciting a response was 3%-17% of concentrations within heads of P. compressa in nature. When moving freely in filtered seawater, competent larvae swam along straight paths in all directions at approximately 0.2 cm s(-1), whereas in water conditioned by P. compressa, most ceased swimming and sank at approximately 0.1 cm s(-1). The ability of larvae to rapidly respond (by sinking) to brief encounters with dissolved settlement cues can enhance their rapid transport to the substratum, even in wave-driven turbulent flow.     

Settlement behavior of cyphonautes larvae of the bryozoan Membranipora membranacea in response to two algal substrata

Abstract. Settlement is an important process in the biphasic life histories of many marine invertebrates. Little is known regarding the fine-scale behavioral mechanisms for finding and attaching to a suitable substratum, particularly under conditions that may impose a potential challenge, such as flow. In this study, we examined the settlement behavior of cyphonautes larvae of the bryozoan, Membranipora membranacea, in response to two different algal substrata. Larvae showed a strong preference for settling on the kelp Nereocystis luetkeana over the red alga Mazzaella splendens. We then tested whether the behavioral mechanisms used by larvae to attach to these algae differed when presented with the challenge of flowing water during settlement. We found that larvae exhibited different behaviors on the two species of algae in flowing water. Larvae were more often observed in direct contact with the preferred alga (N. luetkeana) exhibiting fine-scale active search behaviors. On the less preferred alga (M. splendens), larvae were less frequently observed in direct contact with the alga, and appeared to be exhibiting broad-scale passive search behaviors along the surface of the blade. Our results suggest that cyphonautes larvae alter their behavior in response to their preferred settlement habitat.     

Larval settlement of the nemertean egg predator Carcinonemertes errans on the Dungeness crab,Metacarcinus magister

Due to their habitat specificity, marine parasites present excellent systems for studying the processes and patterns of larval settlement. Settlement of Carcinonermertes errans, an egg predator of the Dungeness crab, is described here for the first time. Upon contact with a host individual, competent larvae of C. errans settled on the crab's exoskeleton and migrated under the abdominal flap within 24 h. When removed from the host, recently settled worms retained their larval characteristics. After 48 h on the host, however, metamorphosis proceeded and larvae became juvenile worms. Additional field studies showed that competent larvae were present in the waters of the Coos Bay Estuary during the months of August through early November, could infect crab hosts directly from the water column, and exhibited density‐dependent gregarious settlement.     

Host-finding behaviour and navigation capabilities of symbiotic zooxanthellae

Past studies have shown that the initiation of symbiosis between the Red-Sea soft coral Heteroxenia fuscescens and its symbiotic dinoflagellates occurs due to the chemical attraction of the motile algal cells to substances emanating from the coral polyps. However, the resulting swimming patterns of zooxanthellae have not been previously studied. This work examined algal swimming behaviour, host location and navigation capabilities under four conditions: (1) still water, (2) in still water with waterborne host attractants, (3) in flowing water, and (4) in flow with host attractants. Algae were capable of actively and effectively locating their host in still water as well as in flow. When in water containing host attractants, swimming became slower, motion patterns straighter and the direction of motion was mainly towards the host—even if this meant advancing upstream against flow velocities of up to 0.5 mm s⁻¹. Coral-algae encounter probability decreased the further downstream of the host algae were located, probably due to diffusion of the chemical signal. The results show how the chemoreceptive zooxanthellae modify their swimming pattern, direction, velocity, circuity and turning rate to accommodate efficient navigation in changing environmental conditions.     

Larval settlement in benthic environments: the effects of velocity and bed element geometry

1. Fluid‐mediated transport can play a key role in determining patterns of distribution and abundance for many benthic invertebrates. One critical challenge in understanding this process is to determine how flow patterns affect larval settlement, especially in those benthic environments where near‐bed flows interact with irregular bed topography to create complex variations in habitat suitability and settlement probability. 2. Boundary‐layer separation over topographical projections on an irregular bed can create two distinct regions of near‐bed flow (i.e., accelerating flow over the forebody and a zone dominated by slower eddies over the aftbody) that may have different effects on larval settlement. 3. We manipulated the flow over a convex roughness element (i.e., hemicylinder) in a flume and examined how the settlement of larvae of the black fly Simulium tribulatum varied with changes in near‐bed velocity and location over the substrate. Larval settlement rate was standardised to correct for variations in larval supply (i.e., among‐trial differences in the concentration of larvae in suspension). 4. Our analyses showed that position on the hemicylinder and near‐bed velocity both affected settlement rate, with a strong interaction effect. In particular, the observed relationship between settlement rate and velocity was negative on the substrate’s forebody and positive on the aftbody. We explore these results by considering potential physical and behavioural mechanisms affecting larval settlement. 5. The presence of a positive relationship between flow and settlement rate in the aftbody may allow settlement on bed elements in habitat where preferred fast‐flow conditions are present, but where settlement would otherwise by hydrodynamically limited. Thus, greater attention to settlement mechanisms in more realistic, topographically complex environments can not only help explain distribution patterns within substrates, but also among substrates and across habitats.     

Larval behaviour and choice of settlement site: Correlation with environmental distribution pattern in an erect bryozoan

Summary This paper concerns the preferred environmental conditions for settlement and metamorphosis of larvae of the erect, rigidly calcified cheilostome bryozoanSchizotheca serratimargo and compares the settlement preferences with in situ distribution of adult colonies. A review of bryozoan larval behaviour and responses to environmental parameters is given in order better to understand larval settlement and metamorphosis in this species. Release of larvae ofS. serratimargo (Hincks) was induced so that each travelled down a path with a limited set of microenvironments in which it could choose to settle. Four different flow velocities, three different depths below the water surface, two different light intensities, three different orientations of plexiglas surfaces, and subdivision of distance of travel into three segments, yielded 216 microenvironments. 466 larvae successfully settled and metamorphosed within the experimental chamber. They metamorphosed in significantly higher numbers in microenvironments with low flow velocity, in the shallowest of the three possible depths, in darker areas, and on nonoverhanging surfaces with little or no sediment. The most important stimulus for settlement was darkness, especially where there was a strong gradient in light intensity.S. serratimargo grows as erect bilaminate bushes on protected flanks of boulders, in small caves, and just within the entrances of larger caves at depths of 5–30 m, as well as on hard debris on open muddy sand floors at 30–35 m depth in the Adriatic and Mediterranean Seas. The experimentally-determined settlement patterns suggest that recruitment is more important than post-settlement interactions or exposure to physical damage in determining local distribution of the species.     

The interplay of substrate nature and biofilm formation in regulating Balanus amphitrite Darwin, 1854 larval settlement

The settlement of marine larvae is influenced by a wide range of physical and biological factors. It is still poorly known how the nature of substrate and the biofilm can interact in regulating settlement patterns of invertebrate larvae. Here we use laboratory experiments focused on settlement behaviour of the barnacle Balanus amphitrite. The aim of this work is to understand whether: (i) the nature of substratum can affect biofilm formation and its structure, (ii) the nature of substratum can affect B. amphitrite larval settlement, (iii) the age of the biofilms and the nature of substrate can interact in influencing larval settlement.Four kinds of substrata (marble, quartz, glass, and cembonit) were biofilmed under laboratory conditions for 5, 10 and 20 days at the temperature of 28 °C. Settlement response was investigated with 5-day-old cyprids. Biofilms were quantitatively and qualitatively analysed by scanning electron microscopy. The settlement of B. amphitrite larvae significantly differed among substrata; also, the patterns of development of biofilm assemblages changed with substrate. In addition, the larval attractiveness of different substrates tends to disappear with biofilm age.     

Predation risk assessment by larval reef fishes during settlement-site selection

Predation rates of marine species are often highest during the transition from the pelagic to the benthic life stage. Consequently, the ability to assess predation risk when selecting a settlement site can be critical to survival. In this study, pairwise choice trials were used to determine whether larvae of three species of anemonefish (Amphiprion melanopus, A. percula and Premnas biaculeatus) are able to (1) assess the predation risk of potential anemone settlement sites through olfactory cues alone and (2) alter their settlement choices depending on the options available (host or non-host anemone). When predation risk was assessed with host and non-host anemone species independently, all species of anemonefish significantly chose the odor associated with the low-risk settlement option over the high-risk site. Most importantly, all species of anemonefish selected water with olfactory cues from their host anemone regardless of predation risk when paired against non-host anemone odor. These results demonstrate that larval reef fishes can use olfactory cues for complex risk assessment during settlement-site selection; however, locating the correct habitat is the most important factor when selecting a settlement site.     

Biofilm history and oxygen availability interact to affect habitat selection in a marine invertebrate

In marine systems, oxygen availability varies at small temporal and spatial scales, such that current oxygen levels may not reflect conditions of the past. Different studies have shown that marine invertebrate larvae can select settlement sites based on local oxygen levels and oxygenation history of the biofilm, but no study has examined the interaction of both. The influence of normoxic and hypoxic water and oxygenation history of biofilms on pre-settlement behavior and settlement of the bryozoan Bugula neritina was tested. Larvae used cues in a hierarchical way: the oxygen levels in the water prime larvae to respond, the response to different biofilms is contingent on oxygen levels in the water. When oxygen levels varied throughout biofilm formation, larvae responded differently depending on the history of the biofilm. It appears that B. neritina larvae integrate cues about current and historical oxygen levels to select the appropriate microhabitat and maximize their fitness.     

Habitat quality and endoparasitism in the Pacific sanddab Citharichthys sordidus from Santa Monica Bay, southern California

Prevalence, mean abundance and mean intensity of the most common endohelminths infecting Pacific sanddab Citharichthys sordidus were compared among six trawl stations in the Santa Monica Bay. These trawl stations varied in their distance from the 8 km effluent of the Hyperion treatment plant which is situated adjacent to Santa Monica Bay. Juvenile stages of Anisakis sp., Corynosoma sp., Lacistorhynchus dollfusi and Tetraphyllidean plerocercoids were found infecting Pacific sanddabs. Significantly higher prevalence and mean intensity of L . dollfusi and Anisakis sp. were seen at the non-outfall stations compared to outfall stations either immediately surrounding or in the vicinity of the outfall. These differences appeared to be related to changes in invertebrate population densities in response to the presence of contaminants. Canonical analysis of discriminance using parasite abundance data discriminated non-outfall from outfall stations. The cestode L . dollfusi explained most of the variability in discriminant scores generated from the analysis and appears to warrant further investigation as a potential bioindicator of pollution exposure in its fish host.     

Larval recruitment of the blue mussel Mytilus edulis: The effect of flow and algae

The mussel Mytilus edulis settlement and distribution was studied on plastic panels with manipulated flow regime (faired, bluff, split and angled) with or without water soluble metabolites of the green alga Cladophora rupestris. The panels were exposed vertically on a device (hydrovane) that ensures their constant orientation in the current during the peak of larval settlement at 1 m depth. In order to investigate larval distribution on the panels, half of them were coated with a silicone vacuum grease that prevents larvae from de-attachment. This grease was not toxic and did not attract or repel larvae. Low densities of larvae on the un-greased plates compared to the greased ones suggested that some of larvae left the substratum. The blue mussel larvae initially settled in regions of reduced shear velocity and then redistribute to the regions of high shear velocity. The presence of the alga increased the density of blue mussel larvae and changed their distribution on the panels. Overall, our results demonstrated that larval recruitment of M. edulis is an active process affected both by boundary-layer hydrodynamics and algal waterborne compounds.     

Effects of near‐bed turbulence on the suspension and settlement of freshwater dreissenid mussel larvae

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Effect of meiofauna on macrofauna recruitment: settlement inhibition of the polychaete Hydroides elegans by the harpacticoid copepod Tisbe japonica

Various aspects of marine macroinvertebrate ecology cannot be understood without detailed knowledge of larval settlement processes. An important effect underscored during the settlement process is the disturbance of marine invertebrate larvae by predators. We demonstrated that biotic disturbance, comprising physical elimination and mortality due to predation and the behavioral irritation of larvae by the harpacticoid copepod Tisbe japonica, prevent a significant portion of larvae of the polychaete Hydroides elegans from settlement on otherwise suitable substrata. Experiments were performed both in the laboratory and the field showing reproducible significant differences in larval settlement and mortality rates between gender-specific copepod treatments and the control. The trend of decreased larval settlement in the presence of copepods coincided with increased larval mortality in these treatments. In the corresponding field experiments, larval settlement and mortality were similar to the ones obtained under laboratory conditions.     

寄主植物对甜菜夜蛾三龄幼虫抗寒力的影响

寄主植物是影响昆虫抗寒性的主要因子之一。研究了不同温度下甜菜夜蛾Spodoptera exigua(Hübner)3龄幼虫取食小白菜、甘蓝、葱和菠菜后,对过冷却能力和体内冷冻保护剂的影响。结果表明,寄主植物对甜菜夜蛾3龄幼虫的过冷却能力有显著性影响,其中以取食甘蓝的幼虫过冷却点最低。温度和寄主植物对其过冷却点、结冰点和虫体含水量有明显的交互作用。寄主植物对其体内的海藻糖含量有显著性影响,而对甘油和糖原含量没有显著性影响。温度和寄主植物仅对海藻糖含量有显著的交互作用。随着温度的升高,取食不同寄主的幼虫体内海藻糖和糖原含量的变化趋势完全相反,认为海藻糖是由糖原转化而来。研究结果提示冬季合适的寄主植物有利于甜菜夜蛾低龄幼虫越冬。     

Hydrodynamic interactions with coral topography and its impact on larval settlement

Coral Reefs - Many benthic larvae rely on ambient flow and turbulence for their dispersal to settlement sites. After reaching the seafloor, larvae must prevent predation as well as overcome flow...