Marine Invertebrate Larvae Research Articles

Sniffing Out New Data and Hypotheses on the Form, Function, and Evolution of the Echinopluteus Post-Oral Vibratile Lobe

The performance requirements of ciliary band feeding explain the convoluted forms of many marine invertebrate larvae. Convolutions increase surface area and therefore feeding rates per unit body volume. We review recent advances in morphology, neural development, and behavior at settlement of the echinoid Lytechinus pictus and provide new ultrastructural and expression data on larvae of its congener, L. variegatus. Larvae of the echinometrid Colobocentrotus atratus contain neurons identified by their expression of nitric oxide synthase (NOS), indicating that this character is not unique to Lytechinus. We hypothesize that in some echinoids the convoluted shape of the post-oral vibratile lobe (POVL) covaries with the distribution of identified sensory neurons to enable olfaction during settlement. An analysis of variation in structural elaboration of the post-oral transverse ciliary band (PTB) within Echinoida and in feeding larvae of other echinoderm classes indicates that only echinoids, but not all echinoids, possess this novel character; larvae that do are distributed heterogeneously within the class. In recognition of this specialized function for the POVL and surrounding ectoderm, and because it is lobate and grows toward the mouth, we propose naming this structure the adoral lobe.

About the Cover.

Next article FreeAbout the CoverPDFPDF PLUSFull Text Add to favoritesDownload CitationTrack CitationsPermissionsReprints Share onFacebookTwitterLinked InRedditEmailQR Code SectionsMoreCover“Biology of Marine Invertebrate Larvae,” the “virtual symposium” in this issue, offers a glimpse of the broad extent of current research on early life stages of marine invertebrates. The images on the cover are a small sample of some of the early developmental stages and larvae that are considered in research, review, and position papers presented in this issue. All stages are about 1 millimeter, or less, in size. Clockwise from top: the large and yolky egg of the sea star Nepanthia troughtoni; a zoea larva of a crab in the family Cancridae; a feeding, late-stage pluteus larva of the sea urchin Tripneustes gratilla; a nonfeeding brachiolaria larva of a sea star collected with a plankton net; a feeding veliger larva of a gastropod in the Thaidae, a family of carnivorous snails; and a nonfeeding, coronate larva of a colonial “moss animal” in the bryozoan genus Bugula. The central image is a confocal z-projection of a phallacidin-labeled planktonic planuliform larva of the nemertean worm Paranemertes peregrina.Larvae occur between the embryonic and juvenile stages of most marine invertebrates. They are small (on the order of 1 mm), slow moving, and usually found suspended in the water where they are dispersed by currents. As individuals they exhibit behavior: they swim, (often) feed, respond to many environmental stimuli, and eventually settle and metamorphose to become juveniles. Larvae are studied for many reasons, including their unusual anatomy and behavior (compared to that of adults), their dynamic development as they change from embryos to juveniles, their dispersal which links populations genetically, and their recruitment as they contribute new individuals to local populations and influence population dynamics. Larval forms and variations among them have also been foci in considerations of animal evolution and animal ecology, and more recently, of how developmental mechanisms and evolutionary forces interact to create the diversity of organisms found among invertebrate phyla. Photo credits: Confocal image of the planuliform larva of Paranemertes peregrina, Svetlana Maslakova (University of Oregon). All other cover images, Richard Emlet (University of Oregon). Cover design: Beth Liles (Marine Biological Laboratory). Next article DetailsFiguresReferencesCited by The Biological Bulletin Volume 216, Number 3June 2009 Published in association with the Marine Biological Laboratory Article DOIhttps://doi.org/10.1086/BBLv216n3cover Views: 43 © 2009 by Marine Biological Laboratory. All rights reserved.PDF download Crossref reports no articles citing this article.

Measurement of the attachment strength of brachiolaria larvae and metamorphic individuals of the sea star Asterina gibbosa by a centrifugation method

Two methods are generally used to measure the adhesive strength of invertebrate larvae: direct measurement with a force transducer connected to the organisms and indirect measurement with a water flow used to dislodge the organisms. Each of these methods, however, has its drawbacks. The present study aimed to design a simple and straightforward method to measure the adhesion strength of marine invertebrate larvae based on centrifugation. This centrifuge technique works in immersed conditions and applies forces acting at 45° to the substratum, therefore mimicking natural conditions. It was tested with three different substrata on two developmental stages of the sea star Asterina gibbosa: the brachiolaria larvae, which use temporary adhesion, and the metamorphic individuals which use permanent adhesion. Measurements were completed by SEM and TEM observations of the larval adhesive organs. The critical detachment force (force required to detach 50% of the larvae) of brachiolaria larvae attached to glass (36 ±9 µN) and rough PMMA (43 ± 16 µN) were equivalent and both significantly higher than the critical detachment force measured on smooth PMMA (11 ± 8 µN). Most metamorphic individuals, on the other hand, resisted to the highest centrifugation speed used, corresponding to a force of 2.13 mN. For the hydrodynamics of larval settlement and metamorphosis, force is the ecologically relevant factor, and adhesion forces obtained by centrifugation are strikingly similar to forces measured for other marine invertebrate larvae with other methods. This indicates the usefulness of the centrifugation technique to compare adhesion of larvae between different species or development stages, or between different treatments.

Adaptation capacity of the larvae of marine invertebrates under variation of environmental parameters as a sensitive test of sea water pollution

The possibility of using the adaptive capacity of the larvae of marine invertebrates under the variation of environmental parameters was studied as a sensitive test of sea water pollution. The object of the research was the larvae of Cryptonatica janthostoma (Naticidae), the environmental factor was the freshening of sea water, and the toxic factor used was copper ions. Adding copper to sea water at a concentration of 0.005 mg/l did not reveal any pronounced effect on the development, behavior and growth of larvae in conditions of normal salinity (32‰), however their adaptive potential decreased with a decrease of salinity to 24‰. The results of the present work and experiments carried out previously enabled us to conclude that before assessing any concentration of pollutant as one not affecting marine organisms, it is necessary to investigate not only their development, growth and survival, but also their adaptive capacity relative to the variation of natural environmental parameters, such as even temperature and salinity.

Serotonergic and FMRFamidergic nervous systems in gymnolaemate bryozoan larvae

A growing body of data from nervous systems of marine invertebrate larvae provides an ideal background for comparisons among higher taxa. The currently available data from Bryozoa, however, do not allow for a consistent hypothesis of an ancestral state for this taxon, which would be necessary for phylogenetic inferences. The larval nervous systems of the four gymnolaemate species Flustrellidra hispida, Bugulafulva, Alcyonidiumgelatinosum, and Bowerbankiagracilis are examined by means of antibody staining against the neurotransmitters serotonin and FMRFamide, as well as against acetylated α-tubulin. Despite considerable variation, a comparison reveals a common pattern of the distribution of serotonin. The neurotransmitter is found in at least two cells in the apical organ as well as in paired axial and lateral nerves emerging from a central nerve nodule. A ring nerve is present below the corona and at least two serotonergic cells are found between the corona cells. Serotonergic coronal cells might represent unique bryozoan features, whereas the remaining elements show resemblance to the situation found in most spiralian taxa. The data do not provide support for a closer relationship of Bryozoa to Phoronida or Brachiopoda.

Molecular detection of native and invasive marine invertebrate larvae present in ballast and open water environmental samples collected in Puget Sound

Non-native marine species have been and continue to be introduced into Puget Sound via several vectors including ship's ballast water. Some non-native species become invasive and negatively impact native species or near shore habitats. We present a new methodology for the development and testing of taxon specific PCR primers designed to assess environmental samples of ocean water for the presence of native and non-native bivalves, crustaceans and algae. The intergenic spacer regions (IGS; ITS1, ITS2 and 5.8S) of the ribosomal DNA were sequenced for adult samples of each taxon studied. We used these data along with those available in Genbank to design taxon and group specific primers and tested their stringency against artificial populations of plasmid constructs containing the entire IGS region for each of the 25 taxa in our study, respectively. Taxon and group specific primer sets were then used to detect the presence or absence of native and non-native planktonic life-history stages (propagules) from environmental samples of ballast water and plankton tow net samples collected in Puget Sound. This methodology provides an inexpensive and efficient way to test the discriminatory ability of taxon specific oligonucleotides (PCR primers) before creating molecular probes or beacons for use in molecular ecological applications such as probe hybridizations or microarray analyses. This work addresses the current need to develop molecular tools capable of diagnosing the presence of planktonic life-history stages from non-native marine species (potential invaders) in ballast water and other environmental samples.

Mechanism of phototaxis in marine zooplankton

The simplest animal eyes are eyespots composed of two cells only: a photoreceptor and a shading pigment cell. They resemble Darwin's 'proto-eyes', considered to be the first eyes to appear in animal evolution. Eyespots cannot form images but enable the animal to sense the direction of light. They are characteristic for the zooplankton larvae of marine invertebrates and are thought to mediate larval swimming towards the light. Phototaxis of invertebrate larvae contributes to the vertical migration of marine plankton, which is thought to represent the biggest biomass transport on Earth. Yet, despite its ecological and evolutionary importance, the mechanism by which eyespots regulate phototaxis is poorly understood. Here we show how simple eyespots in marine zooplankton mediate phototactic swimming, using the marine annelid Platynereis dumerilii as a model. We find that the selective illumination of one eyespot changes the beating of adjacent cilia by direct cholinergic innervation resulting in locally reduced water flow. Computer simulations of larval swimming show that these local effects are sufficient to direct the helical swimming trajectories towards the light. The computer model also shows that axial rotation of the larval body is essential for phototaxis and that helical swimming increases the precision of navigation. These results provide, to our knowledge, the first mechanistic understanding of phototaxis in a marine zooplankton larva and show how simple eyespots regulate it. We propose that the underlying direct coupling of light sensing and ciliary locomotor control was a principal feature of the proto-eye and an important landmark in the evolution of animal eyes.

Egg Energetics, Fertilization Kinetics, and Population Structure in Echinoids With Facultatively Feeding Larvae

Larvae of marine invertebrates either arise from small eggs and feed during their development or arise from large eggs that proceed to metamorphosis sustained only from maternal provisioning. Only a few species are known to possess facultatively feeding larvae. Of about 250 echinoid species with known mode of development, only two, Brisaster latifrons and Clypeaster rosaceus, are known to develop through facultatively planktotrophic larvae. To obtain more information on this form of development and its consequences, we determined egg size and egg energetic and protein content of these two species. We found that eggs of B. latifrons resemble those of species with nonfeeding larvae in these characteristics more than those of C. rosaceus. We also compared DNA sequences of the cytochrome oxidase (COI) gene from the Caribbean C. rosaceus to those of the sympatric planktotrophic developer C. subdepressus and also to those of the eastern Pacific species C. europacificus to estimate the degree of divergence between species with different developmental modes. Comparison of COI sequences of C. rosaceus from Panama and Florida revealed that there is no geographic differentiation in this species. Cross-fertilization experiments between C. rosaceus and C. subdepressus indicated that bidirectional gametic incompatibility has evolved between the two species.

Field-based video observations of wild barnacle cyprid behaviour in response to textural and chemical settlement cues

Many marine invertebrate larvae respond behaviourally to environmental settlement cues, yet behaviours are often only inferred from settlement patterns or are limited to laboratory studies. The behaviour of wild cypris larvae of Semibalanus balanoides L. was filmed on settlement tiles in the field. Tiles were of five different textures with a nested treatment of crude conspecific adult extract (AE). The effects of texture and AE on eleven defined behaviours were analysed. Texture affected the gross and net exploratory distances, velocity, acceleration and time spent exploring. AE attracted more cyprids during the first minute of immersion and increased the time spent on surfaces. Relatively few arrivals that either travel far and fast, or exit the surface rapidly, may indicate a lower chance of settlement. An increase in time spent on a surface may increase the probability of being in contact with the surface when the sign stimulus to settle occurs.

Proteomic analysis of larvae during development, attachment, and metamorphosis in the fouling barnacle, Balanus amphitrite

The barnacle, Balanus amphitrite, is one of the primary model organisms for rocky-shore ecology studies and biofouling research. This barnacle species has a complex life cycle during which the swimming nauplius molts six times and transforms into a cyprid stage. Cyprids must attach to a surface to metamorphose into a juvenile barnacle. To clarify the overall profile of protein expression during larval development and metamorphosis, 2-DE was used to compare the proteome of the nauplius, the swimming cyprid, the attached cyprid, and the metamorphosed cyprid. The proteome of the swimming cyprid was distinctly different from that of other life stages and had about 400 spots. The proteomes of the attached and metamorphosed cyprids were similar with respect to major proteins but had significantly lower numbers of spots compared to that of swimming larval stages. Obviously, synthesis of most proteins from swimming cyprids was switched off after attachment and metamorphosis. Our advanced MS analysis (MALDI-TOF/TOF MS/MS) allowed us to identify the proteins that were differentially and abundantly expressed in the swimming cyprid. These proteins included signal transduction proteins (adenylate cyclase and calmodulin) and juvenile hormone binding proteins. In summary, for the first time, we have analyzed the global protein expression pattern of fouling marine invertebrate larvae during metamorphosis. Our study provides new insights into the mechanisms of barnacle larval metamorphosis and also provides a foundation for exploring novel targets for antifouling treatments.

Evaluation of PCR methods for fixed bivalve larvae

Investigating the spatio-temporal patterns of planktonic larvae is fundamental to studies regarding stock assessment and larval dispersal of commercial and non-commercial, i.e. invasive or rare marine invertebrates. Because of the difficulty involved in morphological identification of marine invertebrate larvae, various molecular methods based on PCR have been used to enhance taxonomic resolution. In previous studies, different methods for the preservation or pretreatment of larvae were applied in each case. However, no comparative studies have been conducted to determine the optimal method for PCR testing for bivalve larvae, and no information is available regarding the selection of an appropriate method.This study compared the PCR success rate of 6 pretreatment methods for larvae of the Mediterranean blue mussel, which was preserved using different fixatives (70% ethanol, 100% ethanol, 70% acetone and 10% formalin). The results revealed that the success rate of PCR was different for each pretreatment; moreover, the use of ammonium sulphate and Tween 20 buffer with proteinase K digestion was found to be the most effective method. Some pretreatments showed lower success rates for long-fixed larvae than for short-fixed larvae for formalin-fixed larvae; however, the success rate of PCR amplification for ethanol-fixed larvae pretreated by this method did not decrease through 1-year fixation. In addition, this pretreatment showed a high success rate for different fixation periods. These findings suggest that the selection of the pretreatment method is critically important for successfully amplifying larval DNA and that the pretreatment involving the use of ammonium sulphate prior to PCR amplification enables the use of fixatives for preserving bivalve larvae. This method will be utilized in various field studies and molecular genetic studies.

A robotic molecular method for in situ detection of marine invertebrate larvae

Knowledge of the temporal and spatial abundance of invertebrate larvae is critical to understanding the dispersal capabilities and recruitment potential of marine and aquatic organisms. Traditional microscopic analyses are time-consuming and difficult given the diversity of larval species and a frequent lack of discriminating morphological characteristics. Here, we describe a sensitive rRNA targeted sandwich hybridization assay (SHA) that uses oligonucleotide probes to detect and enumerate the larvae of invasive green crabs (Carcinus maenas), native blue mussels (Mytilus), native barnacles (Balanus) and polychaetes (Osedax and Ophelia) that occur in the Monterey Bay National Marine Sanctuary, California. Laboratory-based assays demonstrate specificity, high sensitivity, and a quantitative response to cultured samples from three of the target organisms. Oligonucleotide probes were then printed in arrays on nitrocellulose membranes and deployed in our robotic Environmental Sample Processor (ESP) to detect larvae in situ and autonomously. We demonstrate that the SHA-detection method and ESP robot can be used for near real-time, in situ detection of larval species in the marine environment.

Go with the flow: tidal import and export of larvae from semi-enclosed bays

There is a growing body of evidence that suggests the effective functioning of marine reserves is dependent on the dispersal and recruitment of larvae. Enhanced production inside reserves is predicted to lead to a net larval export and increased settlement and recruitment outside reserve boundaries. However, larval retention in bays is also well documented. Since bays are increasingly being used as reserve areas, planktonic larvae of benthic marine invertebrates were sampled from two semi-enclosed marine reserves during flood and ebb tides to determine whether these bays are acting as net exporters of larvae. Neither reserve was a net importer or exporter of species richness, larval abundance or diversity, although one reserve showed a small export of species richness during the hours of darkness. Both reserves balanced the net import of some species with a net export of others, which was generally related to adult or larval abundance, although exceptions were found in one reserve. Significant effects of light were found, with the net import or export of some species occurring exclusively during either the hours of daylight or darkness. An increased understanding of larval sink-source dynamics in bays is essential for ensuring their effective use as marine reserves to meet specific conservation needs.

Size-Specific Predation on Marine Invertebrate Larvae

Predation on planktonic larval stages is frequently a major source of mortality for the offspring of benthic marine invertebrates. Mortality rate likely varies with larval size and developmental stage, but few experiments have measured how these factors affect predation rates. I used experimental reductions in egg size to test how variation in larval size affects the likelihood of predation during planktonic development. Blastomeres of the sand dollar Dendraster excentricus were separated at the two-cell stage to produce half-sized zygotes. Larvae resulting from this manipulation were tested for their susceptibility to predation relative to whole-sized siblings at four ages. Individuals from each size class were simultaneously presented as prey items to five predators (crab zoeae, crab megalopae, chaetognaths, solitary tunicates, and postlarval fish) in the laboratory. Four predators consumed significantly more half-sized larvae than whole-sized larvae, but one predator type (postlarval fish) consumed more whole-sized larvae. Predators that consumed more half-sized larvae also preferentially consumed younger larvae. In contrast, postlarval fish showed no significant prey preference based on larval age. These results suggest that assumptions of constant mortality rates during development should be modified to account for the effects of larval size and age.

Interaction of conspecific cues in Balanus amphitrite Darwin (Cirripedia) settlement assays: Continued argument for the single-larva assay

Gregariousness in marine invertebrate larvae is an important regulator of benthic community structure. Previous laboratory settlement assays employing Balanus amphitrite Darwin cyprids found gregarious effects with as few as 3 larvae well−1, together with modulation of such effects by chemical cues. Here, the relationship between settlement rate and larval density was rigorously tested through a fully randomised design. Seawater conditioned with adult B.amphitrite was tested alongside unconditioned seawater to determine the effect of a conspecific cue on gregarious interactions. Gregarious effects were detected in both conditioned and unconditioned seawater at ≤4 larvae well−1. In untreated seawater, settlement rate increased linearly with larval density, levelling off at densities of ≥10 larvae well−1. In conditioned seawater, settlement induction was observed at ≤4 larvae well−1, switching to inhibition at 6, 8 and 10 larvae well−1, before asymptoting at the highest densities tested. These results advocate the use of individual larvae in laboratory assays that investigate factors stimulating barnacle settlement.

A sampling device to quantify offspring release of sessile marine invertebrates

Summary Quantifying the rate of propagule release is of most importance to estimate reproductive output of natural populations, but simple methods to obtain such data are seldom reported. We designed and tested an inexpensive apparatus capable of reliably measure the release of gametes, eggs or larvae of sessile marine invertebrates in vertical walls. A population of the acorn barnacle Chthamalus bisinuatus was sampled with this trap over 68d to obtain a time series of naupliar release. An apparent semilunar trend is shown, indicating the effectiveness of this sampling method.

Approach toward species identification and quantification of marine invertebrate larvae by DNA analysis (<Feature> Ecological perspectives for the DNA barcoding project)

Approach toward species identification and quantification of marine invertebrate larvae by DNA analysis (<Feature> Ecological perspectives for the DNA barcoding project)

Co‐option and dissociation in larval origins and evolution: the sea urchin larval gut

The origin of marine invertebrate larvae has been an area of controversy in developmental evolution for over a century. Here, we address the question of whether a pelagic "larval" or benthic "adult" morphology originated first in metazoan lineages by testing the hypothesis that particular gene co-option patterns will be associated with the origin of feeding, indirect developing larval forms. Empirical evidence bearing on this hypothesis is derivable from gene expression studies of the sea urchin larval gut of two closely related but differently developing congenerics, Heliocidaris tuberculata (feeding indirect-developing larva) and H. erythrogramma (nonfeeding direct developer), given two subsidiary hypotheses. (1) If larval gut gene expression in H. tuberculata was co-opted from an ancestral adult expression pattern, then the gut expression pattern will remain in adult H. erythrogramma despite its direct development. (2) Genes expressed in the larval gut of H. tuberculata will not have a coordinated expression pattern in H. erythrogramma larvae due to loss of a functional gut. Five structural genes expressed in the invaginating archenteron of H. tuberculata during gastrulation exhibit substantially different expression patterns in H. erythrogramma with only one remaining endoderm specific. Expression of these genes in the adult of H. erythrogramma and larval gut of H. tuberculata, but not in H. erythrogramma larval endoderm, supports the hypothesis that they first played roles in the formation of adult structures and were subsequently recruited into larval ontogeny during the origin and evolution of feeding planktotrophic deuterostome larvae.

Coastline configuration as a determinant of structure in larval assemblages

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 352:67-75 (2007) - DOI: https://doi.org/10.3354/meps07156 Coastline configuration as a determinant of structure in larval assemblages M. Jessopp1,*, O. R. Mulholland2, R. McAllen1, M. P. Johnson3, T. P. Crowe2, A. L. Allcock3 1Department of Zoology, Ecology and Plant Science, University College Cork, Distillery Fields, North Mall, Cork, Ireland 2School of Biology and Environmental Science, University College Dublin, Belfield, Dublin 4, Ireland 3School of Biology and Biochemistry, Queen’s University Belfast, 97 Lisburn Road, Belfast BT9 7BL, UK *Email: mjessopp@gmail.com ABSTRACT: The richness and turnover of coastal larval pools set upper limits for biodiversity in coastal systems. For particular local systems, such as embayments, the characteristics of the local larval pool are determined by the relative contributions of locally produced and external larvae. The balance between these sources partially reflects the extent of tidal exchange and is hence related to system size and flushing time. Larvae of benthic marine invertebrates were sampled from 8 bays along the Irish coast to investigate the effect of coastline configuration on the characteristics of the larval pool. Flushing time explained 34.5% of the variability in species richness from a series of daily samples. Many of the potentially relevant environmental variables are correlated, limiting the potential for individual variables to be examined in isolation. We therefore used a principal components analysis to describe the major patterns in environmental variability across bays. The second principal component separated bays along a gradient of increasing depth, salinity, tidal range and flushing time. Scores along this component were generally better predictors of the larval pool than single variables, explaining as much as 61.2% of the variation in species richness, diversity and similarity between dates. Deeper bays, with more saline water and longer flushing times, tended to have richer and more diverse larval pools, with a greater consistency in species composition between sample dates. No relationship was found between environmental variables and larval abundance. Our results suggest that flushing time, particularly when in combination with topographic variables, chlorophyll, tidal range and salinity, may be a useful predictor for the richness and turnover of local larval pools. KEY WORDS: Flushing time · Larval retention · Larval dispersal · Larval pool Full text in pdf format PreviousNextCite this article as: Jessopp M, Mulholland OR, McAllen R, Johnson MP, Crowe TP, Allcock AL (2007) Coastline configuration as a determinant of structure in larval assemblages. Mar Ecol Prog Ser 352:67-75. https://doi.org/10.3354/meps07156 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 352. Online publication date: December 20, 2007 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2007 Inter-Research.

Settlement of the barnacle Balanus improvisus: The roles of dopamine and serotonin

The regulation of the settlement process in barnacles has attracted research interest due to their role as fouling organisms. The involvement of neurotransmitters in the regulation of settlement of marine invertebrate larvae has been described in several species. In this work, we reported the effects of the neurotransmitters dopamine and serotonin on the settlement of cyprids of the barnacle Balanus improvisus and described differences among cyprids of different ages. Also, we tested the effects of dopaminergic and serotonergic agonists and antagonists on settlement of cyprid larvae. We found that dopamine significantly stimulated settlement of 2‐ and 4‐day‐old cyprids, while serotonin exerted an inhibitory effect, regardless of cyprid age. The agonists and antagonists to the two neurotransmitters were not able to stimulate settlement but in general had either an inhibitory or no effect. We compared our results to those previously reported of the roles of dopamine and serotonin in the settlement of Balanus amphitrite. There appeared to be striking differences in the effects of these neurotransmitters between the two species because it has been reported that serotonin induces settlement in B. amphitrite and that dopamine inhibits it. This suggested that dopamine and serotonin play pivotal roles in settlement of barnacle but may act in different ways in the two species.