Development to metamorphosis of the nemertean pilidium larva

<p>Abstract</p> <p>Background</p> <p>The nemertean pilidium is one of the most notable planktotrophic larval types among marine invertebrates. The juvenile forms inside the larva from a series of isolated rudiments, called the imaginal discs. The development culminates in catastrophic metamorphosis, in which the larval body is consumed by the juvenile worm. Although the pilidium was first described in 1847, and is commonly found among marine plankton, there is not a single complete description of its development. The few published studies of pilidial development are based on observations of typically unidentified larvae opportunistically collected from plankton at various developmental stages.</p> <p>Results</p> <p>The development of <it>Micrura alaskensis</it>, a common Northwest Pacific coast intertidal nemertean, is described from fertilization to metamorphosis. A staging scheme is proposed based on characteristic developmental milestones. Three pairs of imaginal discs develop as invaginations of larval epidermis. The cephalic discs invaginate from the larval epidermis above the ciliated band, while the cerebral organ discs and the trunk discs invaginate below the ciliated band. All paired imaginal disc invaginations are closely associated with different portions of the larval ciliated band. In addition, two unpaired rudiments contribute to the juvenile - the proboscis rudiment and the dorsal rudiment, which do not develop as invaginations. A pair of thick-walled esophageal pouches previously thought to represent nephridial rudiments give rise to the juvenile foregut. Branched rudiments of protonephridia, and their efferent ducts are also described. Larval and juvenile serotonergic nervous systems are briefly described. Development of the juvenile is completed by 5-8 weeks at 11-15 degrees C. During the rapid metamorphosis the juvenile emerges from and devours the larva.</p> <p>Conclusions</p> <p>This study is the first description of pilidial development from fertilization to metamorphosis in a single species. It is illustrated with photomicrographs of live larvae, diagrams, confocal images, and videos. The findings are discussed in the context of previously published accounts of pilidial development, with which they disagree on several accounts. The results described here indicate a different number, origin and fate of various juvenile rudiments. The proposed staging scheme will be useful in subsequent studies of pilidial development.</p

'Shear thickening' in non-shear flows: the effect of microstructure

The bizarre behaviour of a cornstarch suspension (sometimes called oobleck) is well known to all of us who have led public engagement events. At the right solids fraction, it flows smoothly at slow speeds, but can be shattered with a quick spoon movement; if you prepare a large enough sample, you can run across the surface (but if you stand still, you will sink). In rheology circles this phenomenon is known as shear thickening, though the flows described above are not necessarily shear-dominated. In recent years there has been a proliferation of research on the mechanism behind true shear thickening, using both experiments and numerical simulations of shear flows. The understanding of the underlying mechanism is improving markedly. But the paper ‘Microstructure and thickening of dense suspensions under extensional and shear flows’ (Seto, Giusteri & Martinello, J. Fluid Mech., vol. 825, 2017, R3) is the first to consider more general flows. We have, for the first time, simulations of thickening in extensional flows, which are a far better description of oobleck with a runner on top – and can begin to quantify the difference between the idealised shear thickening and the extension thickening that happens in practice

Sampling multiple life stages significantly increases estimates of marine biodiversity

Biodiversity assessments are critical for setting conservation priorities, understanding ecosystem function and establishing a baseline to monitor change. Surveys of marine biodiversity that rely almost entirely on sampling adult organisms underestimate diversity because they tend to be limited to habitat types and individuals that can be easily surveyed. Many marine animals have planktonic larvae that can be sampled from the water column at shallow depths. This life stage often is overlooked in surveys but can be used to relatively rapidly document diversity, especially for the many species that are rare or live cryptically as adults. Using DNA barcode data from samples of nemertean worms collected in three biogeographical regions—Northeastern Pacific, the Caribbean Sea and Eastern Tropical Pacific—we found that most species were collected as either benthic adults or planktonic larvae but seldom in both stages. Randomization tests show that this deficit of operational taxonomic units collected as both adults and larvae is extremely unlikely if larvae and adults were drawn from the same pool of species. This effect persists even in well-studied faunas. These results suggest that sampling planktonic larvae offers access to a different subset of species and thus significantly increases estimates of biodiversity compared to sampling adults alone. Spanish abstract is available in the electronic supplementary material.Fil: Maslakova, Svetlana A.. University of Oregon; Estados UnidosFil: Ellison, Christina I.. University of Oregon; Estados UnidosFil: Hiebert, Terra C.. University of Oregon; Estados UnidosFil: Conable, Frances. University of Oregon; Estados UnidosFil: Heapy, Maureen C.. University of Oregon; Estados UnidosFil: Venera Pontón, Dagoberto E.. Smithsonian Tropical Research Institute; PanamáFil: Norenburg, Jon L.. National Museum Of Natural History. Departamento de Zoología. Area de Invertebrados; Estados UnidosFil: Schwartz, Megan L.. University of Washington; Estados UnidosFil: Boyle, Michael J.. Smithsonian Tropical Research Institute; PanamáFil: Driskell, Amy C.. National Museum Of Natural History. Departamento de Zoología. Area de Invertebrados; Estados UnidosFil: Macdonald, Kenneth S.. National Museum Of Natural History. Departamento de Zoología. Area de Invertebrados; Estados UnidosFil: Zattara, Eduardo Enrique. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; ArgentinaFil: Collin, Rachel. Smithsonian Tropical Research Institute; Panam

Statistical Parsimony Networks and Species Assemblages in Cephalotrichid Nemerteans (Nemertea)

BACKGROUND: It has been suggested that statistical parsimony network analysis could be used to get an indication of species represented in a set of nucleotide data, and the approach has been used to discuss species boundaries in some taxa. METHODOLOGY/PRINCIPAL FINDINGS: Based on 635 base pairs of the mitochondrial protein-coding gene cytochrome c oxidase I (COI), we analyzed 152 nemertean specimens using statistical parsimony network analysis with the connection probability set to 95%. The analysis revealed 15 distinct networks together with seven singletons. Statistical parsimony yielded three networks supporting the species status of Cephalothrix rufifrons, C. major and C. spiralis as they currently have been delineated by morphological characters and geographical location. Many other networks contained haplotypes from nearby geographical locations. Cladistic structure by maximum likelihood analysis overall supported the network analysis, but indicated a false positive result where subnetworks should have been connected into one network/species. This probably is caused by undersampling of the intraspecific haplotype diversity. CONCLUSIONS/SIGNIFICANCE: Statistical parsimony network analysis provides a rapid and useful tool for detecting possible undescribed/cryptic species among cephalotrichid nemerteans based on COI gene. It should be combined with phylogenetic analysis to get indications of false positive results, i.e., subnetworks that would have been connected with more extensive haplotype sampling

DNA barcode sequencing results for bivalve larvae collected on N. Atlantic cruises at cold seeps sites off of Barbados, the East coast of the U.S. and the Gulf of Mexico (SEEPC project)

Dataset: SEEPC Bivalve DNA Barcoding ResultsDNA barcode sequencing results for bivalve larvae collected on N. Atlantic cruises at cold seeps sites off of Barbados, the East coast of the U.S. and the Gulf of Mexico (SEEPC project). This dataset provides the closest GenBank match for larval bivalve specimens resulting from DNA barcode sequencing, the percent match of the identification, and some information on the specimen collections. For a complete list of measurements, refer to the full dataset description in the supplemental file 'Dataset_description.pdf'. The most current version of this dataset is available at: https://www.bco-dmo.org/dataset/688111NSF Division of Ocean Sciences (NSF OCE) OCE-103045

The trochoblasts in the pilidium larva break an ancient spiralian constraint to enable continuous larval growth and maximally indirect development

Abstract Background Nemertean embryos undergo equal spiral cleavage, and prior fate-mapping studies showed that some also exhibit key aspects of spiralian lineage-based fate specification, including specification of the primary trochoblasts, which differentiate early as the core of the prototroch of the spiralian trochophore larva. Yet it remains unclear how the nemertean pilidium larva, a long-lived planktotroph that grows substantially as it builds a juvenile body from isolated rudiments, develops within the constraints of spiral cleavage. Results We marked single cells in embryos of the pilidiophoran Maculaura alaskensis to show that primary, secondary, and accessory trochoblasts, cells that would make the prototroch in conventional spiralian trochophores (1q2, 1q12, and some descendants of 2q), fully account for the pilidium’s primary ciliary band, but without undergoing early cleavage arrest. Instead, the primary ciliary band consists of many small, albeit terminally differentiated, cells. The trochoblasts also give rise to niches of indefinitely proliferative cells (“axils”) that sustain continuous growth of the larval body, including new ciliated band. Several of the imaginal rudiments that form the juvenile body arise from the axils: in particular, we show that cephalic imaginal disks originate from 1a2 and 1b12 and that trunk imaginal disks likely originate from 2d. Conclusions The pilidium exhibits a familiar relation between identified blastomeres and the primary ciliated band, but the manner in which these cells form this organ differs fundamentally from the way equivalent cells construct the trochophore’s prototroch. Also, the establishment, by some progeny of the putative trochoblasts, of indeterminate stem cell populations that give rise to juvenile rudiments, as opposed to an early cleavage arrest, implies a radical alteration in their developmental program. This transition may have been essential to the evolution of a maximally indirect developing larval form—the pilidium—among nemerteans

Additional file 3: Movie 3. of Development of a lecithotrophic pilidium larva illustrates convergent evolution of trochophore-like morphology

Transverse sections (from apical to vegetal) showing anatomy of the “pileus” stage of Micrura sp. “dark.” A running z-projection movie of the confocal z-series used to make Figure 5e1-e3. Specimen stained with phalloidin (white) and propidium iodide (orange). Scale bar 50 µm. (MOV 13778 kb

MOESM1 of The trochoblasts in the pilidium larva break an ancient spiralian constraint to enable continuous larval growth and maximally indirect development

Additional file 1: Fig. S1. Summary of examined labeled domains. Fluorescence ratio confocal images, maximum intensity projected, of labeled domains organized by quadrants (columns) and quartets (rows). These are the same panels shown in Figs. 2, 3, 4 and 5. Apical organ up on all panels. Labeled domains in A and D quadrants are shown from the left side, those of B and C are shown from the right side. Scale bar 50 Οm