A study of the morphology,ultrastructure, and histochemistry of the food of the pediveliger of Ostrea edulis

During the settlement of Ostrea edulis L., the foot of the pediveliger implements temporary attachment and cementing. The morphology of the foot has been re-examined to try to clarify its role during this period. The foot has a highly developed nervous system and musculature, but mainly consists of sub-epidermal gland cells of 9 different types. Cells of the first gland contain acid mucopolysaccharide, and open over the ventral surface of the foot. The second type of gland contains a neutral glycoprotein, and opens at the tip of the foot. The third gland opens mid-ventrally half way down the foot, and largely contains an aromatic protein. The fourth type of gland contains mainly proteoglycan, and the cells open mid-ventrally behind the third gland. The fifth gland contains acid mucopolysaccharide, and the cells open onto the heel of the foot. The sixth, seventh, eighth and ninth glands contain a variety of acid mucopolysaccharides, proteoglycans and possibly a neutral glycoprotein; their cells open into the byssus duct, which discharges at the base of the heel. The grouping of the glands, position of the openings of the cells, and histochemical properties of the secretions suggest that they may be involved in localised and general adhesion of the foot during temporary attachment, as well as in cementing of the larva.     

Development and metamorphosis of two psolid sea cucumbers,Psolus chitonoides andPsolidium bullatum,with a review of reproductive patterns in the family Psolidae (Holothuroidea: Echinodermata)

Reproduction and development of two psolid sea cucumbers in waters of the San Juan Archipelago, Washington, were studied from April 1981 through August 1984. Spawning inPsolus chitonoides Clark andPsolidium bullatum Ohshima in the laboratory and in the field waters of the San Juan Archipelago, Washington, occurred from February through may, with heaviest spawning in April and May. The egg ofPsolus chitonoides was bright red in color and 627µm in diameter;Psolidium bullatum released eggs that were golden yellow in color and 330µm in diameter. Maximum measured fecundities (spawned eggs) forPsolus chitonoides andPsolidium bullatum were 34 700 and 3 074, respectively. Developmental schedules were similar in both species. Early cleavages were equal, radial, and holoblastic; slightly subequal cleavage was common inPsolus chitonoides. The gastrula elongated to form a uniformly ciliated larva and the vestibule first appeared as a ventral crescent-shaped depression, after which the larva decreased in length and formed three posterior ridges that encircled the early doliolaria. Just prior to settlement, there was loss of cilia between the three ridges and on the posterior end of the early doliolaria, leaving three ciliary rings that are characteristic of the dendrochirote doliolaria larva. Vertical swimming ensued, and five primary tentacles protruded through the vestibule, with occasional attachment to the substratum using the primary tentacles. Upon settlement, two primary podia emerged and all external ciliation was lost except among tentacular papillae and on the end of tube feet. Timing of appearance and of loss of ciliary rings suggest that the rings function in vertical swimming and substratum selection behaviors. Metamorphosis involved the following simultaneous events: (1) resorption of the preoral lobe, (2) formation of large, dorsal, calcareous ossicles, (3) flattening of the trivium, and (4) proliferation of ventral tube feet on Ambulacral Radii B and E to form the sole. Reproduction in the family Psolidae is reviewed. The large proportion of brooders in this family appears to be related to small size, the possession of protective armor and sole, and geographic distribution (mainly in Antarctica). Lecithotrophic doliolaria larvae with three ciliary rings are produced by large northern psolids.     

Utilization of a fecal mass as food by the pelagic mysis larva of the penaeid shrimp Solenocera atlantidis

The undetached peritrophic membrane of the mysis shrimp larva (Solenocera atlantidis) provides a substratum on which microbes, organic debris and planktonic organisms accumulate. Behavioral, metabolic and biochemical data indicate that this fecal mass serves as a concentrated food resource for the larva.     

Observations on the behaviour of the pediveliger of Ostrea edulis during attachment and cementing

The behaviour of pediveligers of Ostrea edulis L. during attachment can be divided into 5 clearly defined phases. These behavioural phases are generally successive, and may be a hierarchy of fixed motor patterns which terminates in the consummatory act of cementing. During attachment, the larva utilises two different mechanisms of movement: the first, a fast, smooth, gliding action is probably due to cilia only; the second, a jerky, muscular method which becomes progressively slower as cementing is approached, is related to the burrowing movements of adult lamellibranchs. The byssus thread, which is discharged during the period of muscular locomotion, was the only secretion seen, but the site of its discharge could not be determined by direct observation.     

Abundance,spatial distribution,and size structure of populations of Oreaster reticulatus (Echinodermata: Asteroidea) on sand bottoms

Three populations of Oreaster reticulatus (Linnaeus, 1758) inhabiting shallow-water (<4 m) seagrass habitats in the Grenadines (West Indies, Caribbean Sea) were associated predominantly with beds of Halodule wrightii. Occupation of fringing inshore areas of bare sand was inversely related to wave action; even where sandy patches occurred offshore, the preferred substratum was H. wrightii. The association of O. reticulatus with H. wrightii is related to the asteroid's microphagous feeding habit and the availability of food resources associated with the seagrass. O. reticulatus rarely occurred on dense beds of Thalassia testudinum, but was moderately abundant in areas of sparse cover. Differences in the occurrence of O. reticulatus among seagrass types may be related to factors afdecting foraging effort, such as the tractability of the substratum and mobility upon it. Populations of O. reticulatus exhibited an aggregated dispersion within beds of H. wrightii, possibly attributable to local substratum heterogeneity and/or reproductive behavior. Increased turbulence induced migration to deeper water and markedly increased aggregation along offshore boundaries. The populations were primarily adults, with some late juvenile stages. The paucity of juveniles and their cryptic behavior and coloration suggest that settlement and early postmetamorphic development occurs in alternate habitats, such as dense beds of T. testudinum. Interpopulation differences in size structure may be associated with differences in the quality and availability of food sources.     

Larvae of a colonial ascidian use a non-contact mode of substratum selection on a coral reef

The rate at which larvae successfully recruit into communities of marine benthic invertebrates is partially dependent upon how well larvae avoid benthic predators and settle on appropriate substrata. Therefore, to be able to predict recruitment success, information is needed on how larvae search for settlement sites, whether larvae preferentially settle on certain substrata, and the extent to which there are adequate cues for larvae to find these substrata. This article describes how larvae of the colonial ascidian Diplosoma similis find settlement sites on a coral reef. Direct field observations of larval settlement were made on a fringing reef in Kaneohe Bay, Oahu, Hawaii, between September 1985 and April 1986. A comparison of the substrata that larvae contacted prior to settlement relative to the percentage cover of these substrata on the study reef suggests that larvae are using a non-contact mode of substratum identification to locate suitable settlement sites. This mode of substratum identification allowed 74% of larvae to evade predation by benthic organisms who would otherwise have eaten larvae if they had been contacted. Of those larvae that evaded predation, 88% subsequently settled on the same two substrata upon which most adults are found (dead coral or the green alga Dictyosphaeria cavernosa). This pattern of settlement was probably a result of active selection, since the two substrata cover only 14.4% of the reef's surface and currents had little effect on the direction in which larvae swam. An important contributing factor to the high success rate of larval settlement on suitable substrata was the lack of any temporal decay in substratum preference. It is concluded that for Diplosoma similis larval supply is a sufficient predictor of larval settlement rate. However, for marine invertebrates whose larvae are passively dispersed and exhibit a greater temporal decay in substratum preference, larval settlement should generally have a greater dependency on spatial variation in the abundance of benthic predators and suitable substrata.     

Induction of anti-predator responses in the green-lipped mussel Perna viridis under hypoxia

Hypoxia due to the over enrichment of waters by nutrients is becoming a global problem. In mussels, enhanced byssus thread production is an important adaptation to the presence of crustacean predators and to energetic hydrodynamic regimes. Thread production is an energy-consuming process, so this study used the green mussel Perna viridis (L.) to examine the response to predator exposure combined with hypoxia. Hypoxia is common in sheltered bays in Hong Kong, and the mussels were collected in one such bay, Lok Wo Sha (latitude/longitude: 22o18′ N/114o10′ E) in January, 2009. The predator used in the experiments was the swimming crab Thalamita danae. Oxygen concentrations used in the 48-h experiments ranged from hypoxic to normoxic (1.5 ± 0.3 mg l⁻¹, 3.0 ± 0.3 mg l⁻¹ and 6.0 ± 0.3 mg l⁻¹). Fewer byssus threads which were also shorter and thinner were produced at reduced oxygen levels, no matter if the predator was present or not; the frequency the mussels shed stalks was also lower. Mussels exposed to the predator, however, have enhanced byssus thread production at all oxygen levels when compared with the control. This has highlighted the significance of anti-predator responses for the survival of individuals even under a stressful environment in which energy supply is limited by aerobic metabolism. Interactive effects between oxygen level and predator exposure were observed for the byssus thread production (frequency of shed stalks, mean byssus thread length, cumulative byssus thread volume), with values obtained at 1.5 and 3.0 mg O₂ l⁻¹ being statistically indistinguishable for the control group without predator but not for the predator group. The lack of differences in the byssus thread production at lower oxygen levels in the absence of predator may indicate the minimum amount of byssus that is required for settlement on a substrate.     

Hydrocarbons in the antennal gland secretion of female European beewolves, Philanthus triangulum (Hymenoptera, Crabronidae)

The antennae of female European beewolves (Philanthus triangulum, Hymenoptera: Crabronidae) contain unique gland reservoirs that are inhabited by the symbiotic bacterium ‘Candidatus Streptomyces philanthi’. The reservoir content is secreted into the subterranean brood cells where it serves a dual function: (1) It provides the larvae with spatial information about the orientation of cocoon spinning, and (2) it supplies the larva with the symbiotic bacteria that are needed for the protection of the cocoon against pathogenic microorganisms. We analyzed the hydrocarbon composition of the antennal gland secretion (AGS) by solid phase micro-extraction–gas chromatography–mass spectrometry. We detected 28 substances, the majority of which were saturated and unsaturated hydrocarbons, with minor amounts of some branched alkanes, two ketones, and three unknown substances. There was a strong dimorphism in the chemical composition of the AGS among beewolf females. The two morphs had either (Z)-9-pentacosene (C₂₅-type) or (Z)-9-heptacosene (C₂₇-type) as the main component, and they differed significantly in the relative amounts of most of the other AGS substances. The hydrocarbon composition of the AGS is very similar to that of the postpharyngeal gland of female beewolves, suggesting that the hydrocarbons are universally used for different purposes and transported into the gland from the hemolymph. The putative function of the hydrocarbons in the AGS is discussed.     

Inhibition of larval settlement to a soft bottom benthic community by drifting algal mats: An experimental test

The hypothesis that drifting red algal mats inhibit settlement of planktonic larvae was tested in a field experiment in 1986. Substratum free of algae (caged boxes) was compared with substratum covered with algae (natural substratum and open boxes). Whereas settling densities of 1500 to 5500 ind. m^-2 of the bivalves Macoma balthica, Cardium glaucum and Mya arenaria were observed in sediment without algal cover, no recruits of the same species were found beneath the algae during the period of peak settlement (June to July). The same difference was observed for the polychaete Nereis diversicolor, although in this case occasional individuals were found beneath the algae. The results demonstrate that algae mats may be efficient inhibitors of larval settlement to benthic soft-substratum communities. It is concluded that algae act as a larval filter.     

Scent marking pheromones in lizards: cholesterol and long chain alcohols elicit avoidance and aggression in male Acanthodactylus boskianus (Squamata: Lacertidae)

Femoral gland secretions are believed to play an important role in chemical communication and social organization of lizards. In spite of this, few studies have investigated the chemical composition and the behavioural roles of these secretions. The lacertid lizard Acanthodactylus boskianus is a good example, having these well-developed glands in both sexes. We used GC–MS chemical analysis of gland secretions and y-maze choice test bioassays to investigate the ability of the lizards to detect and respond to different synthetic blends made from compounds identified in the gland secretions. Based upon the GC–MS quantification data, we selected representatives of the main chemical groups (steroids, alcohols, acids, alkanes) detected in the lizard secretions and used these in a behavioural bioassay against controls. Males showed significant avoidance behaviour for cholesterol and alcohol blends, combined with agonistic behaviour towards these stimuli. Females did not show any significant selection to particular odour combinations. The data support the hypotheses that lizards can potentially use femoral gland secretions in chemical odour trails and utilize scent to mark territories and potentially also to establish dominance hierarchies. Cholesterol and long chain alcohols are suggested as potential candidates functioning as scent marking pheromones in A. boskianus.     

Development,settling behaviour,metamorphosis and pentacrinoid feeding and growth of the feather star Florometra serratissima

Embryonic development of the northeastern Pacific feather star Florometra serratissima takes place within a ridged fertilization membrane. Cleavage is radial, resulting in a coeloblastula, and gastrulations is by invagination. Cilia are swollen terminally during ciliogenesis whereas fully grown cilia possess several swellings along the length of their shafts. Young doliolaria larvae begin to hatch from the fertilization membranes 35 h after fertilization (9.5° to 11.5°C); by 4 d the doliolaria has acquired ciliated bands, a vestibular invagination and an antero-ventral adhesive pit. The surface of the larva is covered with a delicate glycocalyx supported by microvilli. Larvae swim along a vertical sinusoidal path just below the water surface; they begin to explore the substratum at 4.5 d and settlement begins as early as 4.6 d, but can be delayed for up to 9 more days. Larvae settle gregariously in culture and it is suggested that gregarious settlement plays a role in the formation and maintenance of adult aggregations of F. serratissima. Metamorphosis into a stalked cystidean following settlement is rapid. Major changes at this period include: loss of cilia; withdrawal of ectoderm from the glycocalyx; covering over of the vestibular invagination; and a 90 degree rotation of the vestibule to the former posterior end of the doliolaria. Transformation from cystidean to pentacrinoid includes the opening of the 5 oral plates, the extension of the 15 papillate tube feet and further elongation of the stalk. The pentacrinoid is able to feed on small food particles. Rudiments of all 10 adult arms are present by 4 months; at 6 months the pentacrinoid has an arm span of 6.5 mm but cirri and pinnules are not yet present.     

Colony specificity of Dufour gland secretions in a functionally monogynous ant

Summary. Queens in colonies of the small myrmicine ant, Leptothorax gredleri Mayr 1855 (Hymenoptera, Formicidae) engage in dominance interactions and form social hierarchies, in which typically only the top-ranking queen lays eggs. Occasionally, queen antagonism escalates to violent mandible fighting, during which the sting is used to apply Dufour gland secretions onto the cuticle of the opponent. Contaminated queens often are attacked by nestmate workers. Here we show that the chemical composition of the Dufour gland is colony-specific and that workers can discriminate between secretions from their own and other colonies. Our findings suggest that Dufour gland secretions are involved in the establishment of hierarchies within a colony. When invading an alien colony the queen presumably employs the secretions during the expulsion of the resident queen. Apparently, Dufour gland secretions play a role in intraspecific queen competition similar to that in slave-making and inquiline formicoxenine ants, where they function as "propaganda substances" in an interspecific context. Received 7 July 1998; accepted 15 September 1998.     

Développement larvaire de deux Tellinacea,Scrobicularia plana (Semelidae) et Donax vittatus (Donacidae)

Scrobicularia plana Da Costa and Donax vittatus L. were reared in the laboratory through settlement. Fertilizable eggs were obtained by perfusing the ovary with 5% 0.1 M ammonium hydroxyde in sea water. S. plana eggs have a thick chorion, inside which the early larval stages develop; they hatch as straight-hinge larvae more than 60 h after fertilization. This brood protection is considered to be an adaptation to osmotic pressure changes and pollution in the environment. D. vittatus eggs have a very thin chorion and are unprotected. Further development is planktotrophic and very similar for the two species. Under laboratory conditions, the pediveliger stage is attained 3 weeks after fertilization and settlement occurs 1 week thereafter. S. plana spat stop growing until a suitable substratum is available. Meanwhile they undergo a byssus drifting important for postlarval dispersion. Post larvae resume growth as soon as a small quantity of fine sand is added to the rearing jar. The exhalant siphon is developed first, when the post larvae reach a length of 600 m; the inhalant siphon is formed later, at a length of approximately 900 m. Evolution from the larval hinge to the juvenile hinge stage occurs sooner in S. plana than in D. vittatus. Comparison of laboratory larval development with field development indicates that spawning occurs in June and August for S. plana in North Wales (UK).     

Settlement and recruitment of sea urchins (Strongylocentrotus spp.) in a sea-urchin barren ground and a kelp bed: are populations regulated by settlement or post-settlement processes?

I sampled recruitment of very small sea urchins (Strongylocentrotus spp.) by using the anesthetic magnesium chloride to remove individuals from substrata collected in sea-urchin barren grounds (barrens) and kelp beds at Naples Reef near Santa Barbara, California, USA. Preliminary sampling found low numbers of newly settled individuals(<0.6 mm test diam) from April–July in 1984 and 1985, and in April, 1986. In early May, 1986, I found many newly settled seaurchins (0.3 to 0.6 mm, 5 to 17 d old), and I compared the densities of the cohort on several types of natural substrata in barrens and kelp-bed habitats. Newly settled individuals of both purple sea urchins (S. purpuratus) and red sea urchins (S. franciscanus) were present in similar, high densities (1 000 S. purpuratus m^-2) on foliose red algal turf, a dominant substratum ofthe kelp bed, and on crustose coralline algae, the dominant substratum of an adjacent barrens. Larvae of S. purpuratus reared and tested in the laboratory showed high rates of settlement on both red algal turf and on crustose coralline algae, but significantly lower rates on rock. Larvae also settled in response to a partiallypurified extract of coralline algae. The reduced settlement on natural rock surfaces relative to either algal treatment and the significant settlement in response to the extract of coralline algae indicate that larvae discriminate between natural substrata and probably respond to a settlement cue other than, or in addition to, a simple microbial (bacterial) film. The similar densities of young recruits of S. purpuratus on dominant substrata of barrens and kelp bed show that, at least in this case, differential settlement cannot explain the high densities of sea urchins in the barrens habitat. Movement between barrens and kelp bed is unlikely given the small sizes of the newly recruited sea urchins relative to the large distances often involved. Reduced post-settlement mortality of newly settled individuals in the barrens remains the most likely mechanism leading to the higher densities of sea urchins in barrens relative to kelp-bed habitats.     

Settlement of barnacle larvae: surface structure of the antennular attachment disc by scanning electron microscopy

The surface morphology of the antennular attachment disc of the cypris larva of Balanus balanoides has been investigated by scanning electron microscopy, and its function is considered in the context of settlent behaviour. It is concluded that the attachment organ acts as an adhesive pad for temporary attachment while the larva explores the substratum. The surface of the disc is covered with villi, which could retain an adhesive substance thought to be produced by the antennular glands. The open-ended hair of the axial sense organ on the disc is shown.     

Some observations on gametogenesis,larval development and substratum selection of the sea pen Ptilosarcus guerneyi

Ptilosarcus (Leioptilus) guerneyi (Gray) maintained in the laboratory, were observed to spawn in late March, 1972. Gametes, developed in the leaf proper, discharged through the mouths of feeding polyps and were fertilized externally in the sea water. The sea pen's eggs are 500 to 600 in diameter; a large female is capable of producing over 200,000 eggs in one season. A pear-shaped and free-swimming planula larva developed 4 days after fertilization, at a temperature of 12 °C. The larvae were ready to settle and metamorphose when 7 days old if favorable substratum was available, but would remain as planulae for at least 30 days if kept in glass dishes only. The 30-day-old larvae would metamorphose if a suitable substratum (coarse sand, for example) was presented. The larvae do not feed and, hence, development is lecithotrophic. Studies of histogenesis showed that metamorphosis greatly enhanced the rate of cellular differentiation. The high fecundity, lecithotrophic development, and the ability of substratum selection by the larvae explain the success of this species in maintaining a high-density population in many areas of sandy substratum in the shallow waters of Puget Sound (USA), despite the fact that it is preyed upon by 7 species of predators.     

Importance of physical and biological settlement cues used at different spatial scales by the larvae of Semibalanus balanoides

The settling cues used by larvae of Semibalanus balanoides (L.) were examined at large (1 m), medium (1 mm), and small (<300 m) spatial scales, corresponding roughly to choices made during broad and close exploration and final inspection within the settling zone. The experiments were carried out at two locations in Canada in May and June 1984. Samples of substratum surfaces from above (n=87 sites unselected by cyprids) and below (n=84 sites selected by cyprids) the upper limit of the barnacle zone were characterized according to 13 quantitative and semi-quantitative variables (diatom cover, macroalgae, detritus) using scanning electron microscopy (SEM). The data were analysed using a stepwise logistic regression. At the large scale, the presence of an alga (Urospora wormskjoldii Mertens) in the upper uncolonized zone and a diatom (Achnantes parvula Kützing) within the settling zone are potential settlement cues. At the medium scale in the settling zone, larvae select clean sites, devoid of detritus and diatoms or on which detrital matter, when present, is of finer texture than on the unchosen sites within the settling zone. At the smallest scale (<300 m, equal to the average maximum width of the larvae), we tested the hypothesis that the larva seeks optimal microheterogeneity (presumably providing good adhesion). Microheterogeneity of the surface was measured immediately under the larva, around the settling organs, the antennules. Our results show that such surface microheterogeneity is significantly greater and the vertical center of gravity of the surface significantly lower on the sites selected by the cyprids than on the unselected contiguous sites. Fractal measurements carried out on selected and contiguous unselected surfaces indicate that larvae discriminate microheterogeneity levels below a step length of 35 m, a value approximately the size of the diameter of the antennular discs (40 m), which are used to explore the substratum and attach.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)     

Substratum relationships of some tropical Pacific intertidal gastropods

Among tropical Indo-West Pacific herbivorous intertidal gastropods, high intertidal neritids, and limpets and certain lower intertidal limpets are restricted to a narrow range of substratum types. Littorinids, planaxids, low intertidal neritids, most lower intertidal limpets, and the trochid Monodonta labio tolerate a wide variety of substrata. This pattern of substratum specialization is related to the overall adaptive strategies of the various groups: those with a large area of soft tissue in contact with the substratum (neritids and limpets) often specialize to a particular substratum, especially at high shore levels, while those in which the area of contact between animal and substratum is small do not. High intertidal neritids and limpets able to live on a wide range of substrata tend to be species of limited geographic distribution. It is concluded that, even within a tropically similar group of animals (raspers in this case) relevant dimensions of niche separation depend upon the structure of the environment as well as upon the structure of the animals themselves.     

Induction of metamorphosis in larvae of the oyster Crassostrea gigas using neuroactive compounds

Chemical (neuroactive compounds at different concentrations and exposure times) and physical (water agitation, light) factors with potential effects on the metamorphosis of larvae of the oyster Crassostrea gigas (Thunberg) larvae have been studied. The neurotransmitters l-dihydroxyphenylalanine (DO), epinephrine (EP), norepinephrine (NE), and acetylcholine (AC) have been identified as very active inducers of metamorphosis, whilst serotonin (SE), dopamine (DA) and potassium (K) were less effective inducers. The -aminobutyric acid (GA) and ammonium (AM) were found ineffective at the concentrations tested. Exposure to 10^-4 M EP for 15 min was sufficient to promote >80% metamorphosis within 48 h, whereas NE required 2 h to exert comparable induction. Maximum induction by DO (>50%) was achieved after 2 h exposure to 10^-4 M. However, unlike EP and NE, DO was lethal at that concentration in the long term. Maximum induction by AC (30% metamorphosis) was achieved at a concentration of 10^-4 M. In contrast to other neurotransmitters, AC induced settlement behaviour, cementation and eventual metamorphosis, yielding postlarvae which were all attached to the substratum. EP and NE triggered the morphogenetic changes, by-passing settlement and leading to postlarvae not cemented to the substratum. DO induced mostly attached spat at low concentrations (10^-5 M) and unattached spat at high concentrations (10^-4 M), and a similar pattern was apparent for the weaker inducers SE and DA. Regarding physical factors, a highly reflectant surface significantly increased the percentage of attached spat obtained, compared to a dark bottom. No consistent effect of water current or light was detected on the production of unattached spat. The three different forms of induction are discussed in relation to different regulatory pathways of settlement and metamorphosis.