The apical sensory organ of a gastropod veliger is a receptor for settlement cues

On the basis of anatomy and larval behavior, the apical sensory organ (ASO) of gastropod veliger larvae has been implicated as the site of perception of cues for settlement and metamorphosis. Until now, there have been no experimental data to support this hypothesis. In this study, cells in the ASO of veliger larvae of the tropical nudibranch Phestilla sibogae were stained with the styryl vital dye DASPEI and then irradiated with a narrow excitatory light beam on a fluorescence microscope. When its ASO cells were bleached by irradiation for 20 min or longer, an otherwise healthy larva was no longer able to respond to the usual metamorphic cue, a soluble metabolite from a coral prey of the adult nudibranch. The irradiated cells absorbed the dye acridine orange, suggesting that they were dying. When larvae not stained with DASPEI were similarly irradiated, or when stained larvae were irradiated with the light beam focused on other parts of the body, there was no loss of ability to metamorphose. Together these data provide strong support for the hypothesis. Potassium and cesium ions, known to induce metamorphosis in larvae of many marine-invertebrate phyla, continue to induce metamorphosis in larvae that have lost the ability to respond to the coral inducer due to staining and irradiation. These results demonstrate that (1) the ASO-ablated larvae have not lost the ability to metamorphose and (2) the ions do not act only on the metamorphic-signal receptor cells, but at other sites downstream in the metamorphic signal transduction pathway.     

The larval ocelli of Golfingia misakiana (Sipuncula, Golfingiidae) and of a pelagosphera of another unidentified species

 The inverse cerebral ocelli of the pelagosphera larva of Golfingia misakiana and of another unidentified larva are composed of two or three sensory cells and one supportive pigmented cell. The sensory cells bear an array of microvilli as well as a single cilium with poor undulation of its membrane; the photoreceptive organelles are regarded as the rhabdomeric type. A striking feature of these cells is the cores, which extend within the microvilli from the tip into the midregion of the cell. It is suggested that these structures are identical with the submicrovillar cisternae found in the cerebral inverse eyes of larvae of Polychaeta. The findings allow the conclusion that in the pelagosphera of the Sipuncula, contrary to the teleplanic veliger larvae of Gastropoda, a lengthy pelagic cycle is not correlated with the development of a ciliary photoreceptor. Additionally, it is assumed that the pigment cup ocelli in larvae of Sipuncula are homologous with the cerebral inverted pigment cup ocelli of larvae of Polychaeta. Accepted: 19 March 1997     

Larval development of hyolithids

The larval development of hyolithids is inferred to have been closely similar to that of primitive gastropods. A trochophore-like larva produced a swollen initial conch and a convex operculum. The initial conch was either subsphaerical and smooth or fusiform with a pointed apex and growth lines. This is taken to indicate an initial development within egg covers or free-living, respectively. A pelagic veliger phase followed. preceded by hatching in the first case. Veliger shells are similar to adult ones. Metamorphosis (transformation of veliger into benthic adult animal) in both hyolithids and gastropods was accompanied by an increase in mortality.     

Comparative susceptibility of veliger larvae of four bivalve mollusks to a Vibrio alginolyticus strain

The susceptibility of 7 d old veliger larvae of the scallops Argopecten ventricosus and Nodipecten subnodosus, the penshell Atrina maura, and the Pacific oyster Crassostrea gigas to a pathogenic strain of Vibrio alginolyticus was investigated by challenging the larvae with different bacterial concentrations in a semi-static assay. The results indicate that the larvae of the 2 scallop species are more susceptible to the V. alginolyticus strain than those of the oyster and the penshell. Signs of the disease were similar to bacillary necrosis described in previous work. Interspecies differences in susceptibility to pathogens are discussed.     

Neural correlates of settlement in veliger larvae of the gastropod,Crepidula fornicata

Settlement behavior of molluscan veliger larvae prior to metamorphosis requires cessation of swimming, accomplished by arrest of prototrochal cilia on the margin of the velum (the larval swimming organ). Ciliary arrest in larvae of gastropods is mediated by an action potential that occurs synchronously across the velum as a consequence of electrical coupling between the prototrochal ciliated cells. We developed a preparation for extracellular recording of such ciliary arrest spikes from intact swimming and crawling veliger larvae of the caenogastropod Crepidula fornicata, using a fine wire electrode. Ciliary arrest spike rates during bouts of substrate crawling were significantly higher than those recorded during preceding swimming periods in larvae that were competent for metamorphosis, but not in precompetent larvae. Spike rates were similar on clean polystyrene substrates, and on substrates that had been coated with a natural cue for metamorphosis (mucus from conspecific adults). We used immunohistochemical methods to localize neuromodulators that might regulate the function of velar cilia. Labeled terminals for serotonin, FMRFamide, and tyrosine hydroxylase (an enzyme for catecholamine synthesis) were located in positions consistent with modulatory effects on the prototrochal ciliated cells. Prototrochal ciliary arrest spike rates and beat frequencies were measured in isolated velar lobes from competent larvae, which were exposed to serotonin, FMRFamide, and dopamine (10^?5 mol L^?1). Serotonin abolished arrest spiking and increased beat frequency; dopamine also increased beat frequency, and FMRFamide depressed it. Competent larvae tested in a small static water column swam to the top of the column when exposed to serotonin, but occupied lower positions than controls when in the presence of dopamine and FMRFamide. The larval nervous system appears to regulate velar functions that are critical for settlement behavior, and is likely to do so by integrating different sensory modalities in an age‐dependent manner.     

Pelagic larvae of the bivalves of Ussuriysky Bay, Sea of Japan

Based on materials of plankton surveying carried out in May–October 2008, the species composition, distribution, and density of pelagic larvae of bivalves of the Ussuriysky Bay (Peter the Great Bay, Sea of Japan) were studied. Larvae of 66 taxa were revealed and identified. The Bivalvia larvae occurred in the plankton during the entire period of study. Two areas, the inner and open part of the bay, which vary in the composition, concentration, and terms of larva occurrence in the plankton were distinguished. The larvae of Mytilus coruscus, Mactra chinensis, Modiolus kurilensis, Ruditapes philippinarum, Mya uzenensis and Teredo navalis were the most numerous and widespread. It was shown that despite high taxonomic diversity, the density of Bivalvia larvae in the Ussuriysky Bay was an order of magnitude lower, even in terms of their maximum number, than in other studied areas of the Peter the Great Bay.     

A pelagic larva of <Emphasis Type="Italic">Plagiopsetta glossa</Emphasis> (Pleuronectiformes,Samaridae), characterized by a trailing gut and S-shaped pelvic bone: description and phylogenetic considerations

 The pelagic larval stage of a pleuronectiform samarid, Plagiopsetta glossa, is described and illustrated, for the first time, from a 8.4-mm BL postflexion specimen collected off Tosa Bay, southern Japan. The larva is distinctive in having broad, semitransparent pterygiophore zones on the dorsal and anal fins, a trailing gut, flexible S-shaped pelvic bone, and a skin fold under the throat. These characteristics are shared with poecilopsettid larvae, suggesting a close relationship between the two families.     

An alternative mode of larval development inScoloplos armiger (O. F. Müller, 1776) (Polychaeta,Orbiniidae)

During a three-year plankton monitoring at the island of Helgoland (German Bight, North Sea), pelagic larvae ofScoloplos armiger (O. F. Müller, 1776) were observed to occur regularly from February to April. Up to the present day, only benthic development within egg cocoons has been described (Anderson, 1959). Planktonic larvae comprise early, ovoid trochophores up to advanced stages with 11 setigers. The main morphological difference between the two larval types is the more numerous differentiation of lateral cilia by the pelagic larvae. Annual alterations in appearance and abundance suggest that reproduction is correlated with water temperature.     

扁玉螺早期发育的实验观察

在实验室条件下人工孵化扁玉螺(Neverita didyma)的卵块,观察了其胚胎发育和幼虫发育过程.扁玉螺的早期发育属间接发生型,其胚胎发育包括卵裂期、囊胚期、原肠胚、膜内担轮幼虫、膜内面盘幼虫;幼虫发育包括面盘幼虫、后期面盘幼虫和匍匐幼虫;匍匐幼虫经变态后发育为稚螺.在水温25～26℃条件下,受精卵发育至膜内面盘幼虫约需38h,5～6d后面盘幼虫冲破卵膜而孵化.扁玉螺面盘幼虫的显著特点是具有1对眼点和1对平衡囊,面盘呈双叶状;后期面盘幼虫的面盘为4叶,呈蝴蝶状,足发达,幼虫既能浮游,又能爬行.后期面盘幼虫进一步生长发育,逐渐转入匍匐生活.     

Enhanced early detection and enumeration of zebra mussel (Dreissena spp.) veligers using cross-polarized light microscopy

Zooplankton with calcareous skeletons are birefringent under cross-polarized light, and thus this technique can be quite useful, indeed sometimes almost essential, for the detection and enumemeration of these types of organisms in plankton samples. A simple and inexpensive application of this technique is described and illustrated with quantitative examples from research on the veligers of the zebra mussel Dreissena polymorpha (Pallas). The time required to detect veligers in plankton samples was decreased by an order of magnitude; the accuracy of counts was substantially improved (15% more than controls), and the time required for counts was greatly reduced (41% of control times). This technique is especially useful in situations in which veligers are difficult to find or see (e.g., at low densities, in samples cluttered with extraneous organisms or material) or when the investigator is inexperienced with plankton sampling and planktonic organisms. The major limitations are its inability to discriminate among various bivalve species that have planktonic larvae and the similar appearance of ostracods which also have calcareous shells. Expanded use of this technique should (1) increase our ability to use plankton sampling for the early detection of veligers during range expansion and reproductive cycles and (2) permit more accurate estimates of veliger abundance.     

A technique for determining the distribution of pelagic fish larvae

A technique is described for studying small scale distributional patterns of fish larvae in the pelagic region. A plankton net is mounted beneath the bows of a small boat, the contents of the net are emptied continually by a suction pump. This system has advantages over previous techniques as it allows a large number of small discrete samples to be taken over a wide area in a short space of time. Preliminary results showed the population of perch larvae in Llyn Tegid is extremely clumped and the overall abundance of the larvae decreases as samples are taken progressively further away from the south-west end of the lake.     

Survival, growth, settlement and metamorphosis of refrigerated larvae of the mussel Mytilus galloprovincialis Lamarck and their use in settlement and antifouling bioassays

Straight-hinge veliger and pediveliger larvae of the mussel Mytilus galloprovincialis were refrigerated for varying periods for use in bioassays. Straight-hinge veliger larvae grew to the umbo-veliger stage after 2 months in the refrigerator, but no pediveligers were observed during the 3-month refrigeration period. The average survival rate of larvae in the refrigerator was 79% after 1 month, but gradually decreased with the refrigeration period, and was as low as 22% after 3 months. All refrigerated larvae grew to the pediveliger stage in the incubator at 17 degrees C at the same rate as that of the control larvae that were not refrigerated. Settlement and metamorphosis of pediveligers from both refrigerated and control groups were facilitated by microbial film and epinephrine and inhibited by phentolamine. Thus, refrigeration can be used as an effective method of storing larvae of M. galloprovincialis for use in assays to assess candidate settlement inducers and antifouling substances.     

Evolution and development of gastropod larval shell morphology: experimental evidence for mechanical defense and repair

SUMMARY The structural diversity of gastropod veliger larvae offers an instructive counterpoint to the view of larval forms as conservative archetypes. Larval structure, function, and development are fine-tuned for survival in the plankton. Accordingly, the study of larval adaptation provides an important perspective for evolutionary-developmental biology as an integrated science. Patterns of breakage and repair in the field, as well as patterns of breakage in arranged encounters with zooplankton under laboratory conditions, are two powerful sources of data on the adaptive significance of morphological and microsculptural features of the gastropod larval shell. Shells of the planktonic veliger larvae of the caenogastropod Nassarius paupertus [ Gould ] preserve multiple repaired breaks, attributed to unsuccessful zooplankton predators. In culture, larvae isolated from concentrated zooplankton samples rapidly repaired broken apertural margins and restored the "ideal" apertural form, in which an elaborate projection or "beak" covers the head of the swimming veliger. When individuals with repaired apertures were reintroduced to a concentrated mixture of potential zooplankton predators, the repaired margins were rapidly chipped and broken back. The projecting beak of the larval shell is the first line of mechanical defense, covering the larval head and mouth and potentially the most vulnerable part of the shell to breakage. Patterns of mechanical failure show that spiral ridges do reinforce the beak and retard breakage. The capacity for rapid shell repair and regeneration, and the evolution of features that resist or retard mechanical damage, may play a more prominent role than previously thought in enhancing the ability of larvae to survive in the plankton.     

Towards a ground pattern reconstruction of bivalve nervous systems: neurogenesis in the zebra mussel <Emphasis Type="Italic">Dreissena polymorpha</Emphasis>

Bivalvia is a taxon of aquatic mollusks that includes clams, oysters, mussels, and scallops. Within heterodont bivalves, Dreissena polymorpha is a small, mytiliform, freshwater mussel that develops indirectly via a planktotrophic veliger larva. Currently, only a few studies on bivalve neurogenesis are available, impeding the reconstruction of a ground pattern in Bivalvia. In order to inject novel data into this discussion, we describe herein the development of the serotonin-like and α-tubulin-like immunoreactive (lir) neuronal components of D. polymorpha from the early trochophore to the late veliger stage. Neurogenesis starts in the early trochophore stage at the apical pole with the appearance of one flask-shaped serotonin-lir cell. When larvae reach the veliger stage, four flask-shaped serotonin-lir cells are present in the apical organ. At the same time, the anlagen of the cerebral ganglia start to form at the base of the apical organ. From the apical organ, one pair of cerebro-visceral connectives projects posteriorly and connects to a posterior larval sensory organ that contains serotonin- and α-tubulin-like flask-shaped cells. Additional, paired serotonin-lir neurites originate from the apical organ and project into the velum. One unpaired stomatogastric serotonin-lir cell develops ventrally to the stomach at the veliger stage. The low number of serotonin-lir cells in the apical organ of bivalve veligers is shared with larvae of basally branching gastropods and scaphopods and is thus considered a feature of the last common ancestor of Conchifera, while the overall simplicity of the larval neural architecture appears to be a specific trait of Bivalvia.     

Larval ecology and morphology in fossil gastropods

The shell of marine gastropods conserves and reflects early ontogeny, including embryonic and larval stages, to a high degree when compared with other marine invertebrates. Planktotrophic larval development is indicated by a small embryonic shell (size is also related to systematic placement) with little yolk followed by a multiwhorled shell formed by a free‐swimming veliger larva. Basal gastropod clades (e.g. Vetigastropoda) lack planktotrophic larval development. The great majority of Late Palaeozoic and Mesozoic ‘derived’ marine gastropods (Neritimorpha, Caenogastropoda and Heterobranchia) with known protoconch had planktotrophic larval development. Dimensions of internal moulds of protoconchs suggest that planktotrophic larval development was largely absent in the Cambrian and evolved at the Cambrian–Ordovician transition, mainly due to increasing benthic predation. The evolution of planktotrophic larval development offered advantages and opportunities such as more effective dispersal, enhanced gene flow between populations and prevention of inbreeding. Early gastropod larval shells were openly coiled and weakly sculptured. During the Mid‐ and Late Palaeozoic, modern tightly coiled larval shells (commonly with strong sculpture) evolved due to increasing predation pressure in the plankton. The presence of numerous Late Palaeozoic and Triassic gastropod species with planktotrophic larval development suggests sufficient primary production although direct evidence for phytoplankton is scarce in this period. Contrary to previous suggestions, it seems unlikely that the end‐Permian mass extinction selected against species with planktotrophic larval development. The molluscan classes with highest species diversity (Gastropoda and Bivalvia) are those which may have planktotrophic larval development. Extremely high diversity in such groups as Caenogastropoda or eulamellibranch bivalves is the result of high phylogenetic activity and is associated with the presence of planktotrophic veliger larvae in many members of these groups, although causality has not been shown yet. A new gastropod species and genus, Anachronistella peterwagneri, is described from the Late Triassic Cassian Formation; it is the first known Triassic gastropod with an openly coiled larval shell.     

Relating the ontogeny of functional morphology and prey selection with larval mortality in Amphiprion frenatus

Survival during the pelagic larval phase of marine fish is highly variable and is subject to numerous factors. A sharp decline in the number of surviving larvae usually occurs during the transition from endogenous to exogenous feeding known as the first feeding stage in fish larvae. The present study was designed to evaluate the link between functional morphology and prey selection in an attempt to understand how the relationship influences mortality of a marine fish larva, Amphiprion frenatus, through ontogeny. Larvae were reared from hatch to 14 days post hatch (DPH) with one of four diets [rotifers and newly hatched Artemia sp. nauplii (RA); rotifers and wild plankton (RP); rotifers, wild plankton, and newly hatched Artemia nauplii (RPA); wild plankton and newly hatched Artemia nauplii (PA)]. Survival did not differ among diets. Larvae from all diets experienced mass mortality from 1 to 5 DPH followed by decreased mortality from 6 to 14 DPH; individuals fed RA were the exception, exhibiting continuous mortality from 6 to 14 DPH. Larvae consumed progressively larger prey with growth and age, likely due to age related increase in gape. During the mass mortality event, larvae selected small prey items and exhibited few ossified elements. Cessation of mass mortality coincided with consumption of large prey and ossification of key elements of the feeding apparatus. Mass mortality did not appear to be solely influenced by inability to establish first feeding. We hypothesize the interaction of reduced feeding capacities (i.e., complexity of the feeding apparatus) and larval physiology such as digestion or absorption efficiency contributed to the mortality event during the first feeding period. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

The ciliary photoreceptor in the teleplanic veliger larvae of Smaragdia sp. and Strombus sp. (Mollusca,Gastropoda)

The cerebrally innervated eyes of the veliger larvae of Smaragdia sp. and Strombus sp. are composed of a lens, a cornea, and an everse retina. The retina contains two different types of cells, ciliary sensory cells and supportive cells which bear one or two cilia. It is suggested that: (a) the ciliary photoreceptors of these teleplanic veliger larvae are correlated with a long pelagic life in the ocean, which can last up to twelve months, and (b) that structural details of the photoreceptors can change during ontogenesis (ciliary vs rhabdomeric). Furthermore, the cilia of the supportive cells apparently tranport lens material and thus play an important role in lens formation. A decomposition mechanism of pigment granules is examined.Abbreviations bb basal body - bp basal plate - c cilium - cc corneal cell - cm ciliary membranes - cw ciliary whorl - ecm extracellular matrix - gr electron-dense granules - l lens - lb lamellar body - mp membranous pieces - mt microtubules - mv microvilli - n nucleus - oc optic cavity - on optic nerve - pg pigment granule - sc sensory cell - sj septate junction - spc supportive cell - v vesicles     

Larval development and metamorphosis in the marine pulmonate Amphibola crenata (Mollusca: Pulmonata)

The development of the free-swimming veliger of Amphibola is followed from hatching to settlement, and the larval structures compared with those of post-metamorphic juveniles and adult snails. Observations of living specimens and light-microscope sections were combined with scanning electron microscopy to build up a composite picture of veliger structure.
Four stages in the development of veligers are recognized, each being characterized by the appearance of organ systems such as the mantle cavity, larval heart, adult heart and kidney, and larval pallial gland. At or after metamorphosis, the larval systems (heart, kidney and pallial gland) disappear, and the developing adult organs move to the positions characteristic of adult snails.
Organogenesis in Amphibola veligers is compared with that of prosobranch and opisthobranch larvae, and with that of pulmonate larvae with direct development. The closest similarity is seen to be with opisthobranch veligers.     

Eastward and Westward Dispersal Across the Tropical Atlantic Ocean of Larvae Belonging to the Genus Bursa (Prosobranchia,Mesogastropoda, Bursidae)

Larvae of the genus Bursa are commonly found in the South Equatorial Current and the Equatorial Undercurrent of the tropical Atlantic Ocean. Identification of Bursa veliger larvae in the plankton of such tropical waters is made possible by a comparison of larval shells with protoconchs of juvenile and well-preserved adult specimens. The relationship between the average current velocity and the estimated minimum duration of larval development of Bursa thomae suggests that there is sufficient time to account for successful trans-Atlantic transport of its veligers in either direction between West Africa and South America. The dispersal of veliger larvae explains the wide geographical distribution attained by the genus Bursa.     

Hybridisations between Mytilus edulis and Mytilus galloprovincialis and performance of pure species and hybrid veliger larvae at different temperatures

The mussels Mytilus edulis L. and Mytilus galloprovincialis Lamark hybridise naturally in the wild along the Atlantic coast of Europe producing a patchwork of mixed pure species and hybrid populations. Individuals of both species were spawned in the laboratory and were hybridised in a series of reciprocal crosses. After 72 h, the proportion of eggs which developed into larvae (%yield) and the proportion of those larvae which had a normal veliger morphology (%normality) were estimated and compared between pure species and hybrid families. There were no significant differences in %yield or %normality between pure species and hybrids, but significant differences were evident between the offspring from different parents irrespective of whether they were hybrids or pure species. Therefore confirmation of hybrid heterosis in laboratory studies should not be based on a single, or a few reciprocal crosses. Hybrid and pure species veliger larvae were grown for approximately 4 weeks at 10, 14 or 20 °C. In all trials, pure M. galloprovincialis larvae grew significantly faster at 20 °C than either reciprocal hybrid or pure M. edulis larvae. Irrespective of temperature, in general, hybrid larvae grew slower than larvae of either pure species. Increased exposure to planktonic predation due to slow growth can be interpreted as selection against hybrids and this may play a role in the structure and distribution of mixed pure species and hybrid populations.