Par protein localization during the early development of Mnemiopsis leidyi suggests different modes of epithelial organization in the metazoa.

In bilaterians and cnidarians, epithelial cell-polarity is regulated by the interactions between Par proteins, Wnt/PCP signaling pathway, and cell-cell adhesion. Par proteins are highly conserved across Metazoa, including ctenophores. But strikingly, ctenophore genomes lack components of the Wnt/PCP pathway and cell-cell adhesion complexes raising the question if ctenophore cells are polarized by mechanisms involving Par proteins. Here, by using immunohistochemistry and live-cell imaging of specific mRNAs, we describe for the first time the subcellular localization of selected Par proteins in blastomeres and epithelial cells during the embryogenesis of the ctenophore Mnemiopsis leidyi. We show that these proteins distribute differently compared to what has been described for other animals, even though they segregate in a host-specific fashion when expressed in cnidarian embryos. This differential localization might be related to the emergence of different junctional complexes during metazoan evolution.

Cannibalism makes invasive comb jelly, Mnemiopsis leidyi, resilient to unfavourable conditions.

The proliferation of invasive marine species is often explained by a lack of predators and opportunistic life history traits. For the invasive comb jelly Mnemiopsis leidyi, it has remained unclear how this now widely distributed species is able to overcome long periods of low food availability, particularly in their northernmost exotic habitats in Eurasia. Based on both field and laboratory evidence, we show that adult comb jellies in the western Baltic Sea continue building up their nutrient reserves after emptying the prey field through a shift to cannibalizing their own larvae. We argue, that by creating massive late summer blooms, the population can efficiently empty the prey field, outcompete intraguild competitors, and use the bloom events to build nutrient reserves for critical periods of prey scarcity. Our finding that cannibalism makes a species with typical opportunistic traits more resilient to environmental fluctuations is important for devising more effective conservation strategies.

Effect of invasive ctenophores Mnemiopsis leidyi and Beroe ovata on low trophic webs of the Black Sea ecosystem.

The study focuses on the impact of life excretion and mucus released by the "biological pollutants" invasive ctenophore Mnemiopsis leidyi and its predator Beroe ovata on the marine environment and lower trophic levels of the Black Sea ecosystem (bacteria, pico-phytoplankton, nano-autotrophic/heterotrophic flagellates, micro-phytoplankton, chlorophyll a, primary production (PP), micro-zooplankton). The chemical and biological variables were analysed in two sets of lab experiments with natural communities from mesotrophic (Gelendzhik) and eutrophic (Varna) coastal waters. While both species altered the chemical properties of experimental media, exerting structural and functional changes in the low food-web biological compartments, the results showed a stronger effect of B. ovata, most likely related to the measured higher rate of excretion and amount of released mucus. In addition the alterations in the Gelendzhik experiment were more pronounced, indicating that environmental implications on lower food-web are more conspicuous in mesotrophic than in eutrophic coastal waters.

QM/MM simulations provide insight into the mechanism of bioluminescence triggering in ctenophore photoproteins.

Photoproteins are responsible for light emission in a variety of marine ctenophores and coelenterates. The mechanism of light emission in both families occurs via the same reaction. However, the arrangement of amino acid residues surrounding the chromophore, and the catalytic mechanism of light emission is unknown for the ctenophore photoproteins. In this study, we used quantum mechanics/molecular mechanics (QM/MM) and site-directed mutagenesis studies to investigate the details of the catalytic mechanism in berovin, a member of the ctenophore family. In the absence of a crystal structure of the berovin-substrate complex, molecular docking was used to determine the binding mode of the protonated (2-hydroperoxy) and deprotonated (2-peroxy anion) forms of the substrate to berovin. A total of 13 mutants predicted to surround the binding site were targeted by site-directed mutagenesis which revealed their relative importance in substrate binding and catalysis. Molecular dynamics simulations and MM-PBSA (Molecular Mechanics Poisson-Boltzmann/surface area) calculations showed that electrostatic and polar solvation energy are +115.65 and -100.42 kcal/mol in the deprotonated form, respectively. QM/MM calculations and pKa analysis revealed the deprotonated form of substrate is unstable due to the generation of a dioxetane intermediate caused by nucleophilic attack of the substrate peroxy anion at its C3 position. This work also revealed that a hydrogen bonding network formed by a D158- R41-Y204 triad could be responsible for shuttling the proton from the 2- hydroperoxy group of the substrate to bulk solvent.

The onset of regenerative properties in ctenophores.

Ctenophores are a clade of animals that branch off at the base of the animal tree. They have a unique and delicate body plan, and distinct pattern forming mechanisms at different life history stages. They have a stereotyped embryonic cell lineage and are highly 'mosaic' as embryos, but most have amazing capacity to regenerate as adults. Unfortunately, only a handful of ctenophore species have been studied in detail. This review summarizes the key features of the regenerative properties of adults, and the characteristics of the embryological onset of regenerative abilities. The genomes of several ctenophore species have already been sequenced, and these resources set the stage for more detailed cellular and molecular analysis of body plan patterning in this group.

Ctenophores: an evolutionary-developmental perspective.

Ctenophores are non-bilaterian metazoans of uncertain phylogenetic position, some recent studies placing them as sister-group to all other animals whereas others suggest this placement is artefactual and ctenophores are more closely allied with cnidarians and bilaterians, with which they share nerve cells, muscles and gut. Available information about developmental genes and their expression and function in ctenophores is reviewed. These data not only unveil some conserved aspects of molecular developmental mechanisms with other basal metazoan lineages, but also can be expected to enlighten the genomic and molecular bases of the evolution of ctenophore-specific traits, including their unique embryonic development, complex anatomy and high cell type diversity.

Development of neuromuscular organization in the ctenophore Pleurobrachia bachei.

The phylogenetic position of the phylum Ctenophora and the nature of ctenphore nervous systems are highly debated topics in modern evolutionary biology. However, very little is known about the organization of ctenophore neural and muscular systems, and virtually nothing has been reported about their embryogenesis. Here we have characterized the neural and muscular development of the sea gooseberry, Pleurobrachia bachei, starting from the cleavage stages to posthatching larvae. Scanning electron microscopy and immunochemistry were used to describe the formation of the embryonic mouth, tentacles, combs, aboral organ, and putative sensory cells. The muscles started their specification at the end of the first day of Pleurobrachia development. In contrast, neurons appeared 2 days after myogenesis, just before the hatching of fully formed cydippid larvae. The first tubulin-immunoreactive neurons, a small group of four to six cells with neuronal processes, was initially recognized at the aboral pole during the third day of development. Surprisingly, this observed neurogenesis occurred after the emergence of distinct behavioral patterns in the embryos. Thus, the embryonic behavior associated with comb cilia beatings and initial muscle organization does not require morphologically defined neurons and their elongated neurites. This study provides the first description of neuromuscular development in the enigmatic ctenophores and establishes the foundation for future research using emerging genomic tools and resources.

Ctenophores from the Oaxaca coast, including a checklist of species from the Pacific coast of Mexico.

Ctenophores are poorly known in the tropical eastern Pacific, including the southern coast of Mexico. Previous records of ctenophores along the Pacific coast have been provided mainly from northern waters. For the coast of Oaxaca state, their occurrence has only been mentioned before at phylum level. In this paper, we provide the first three records of ctenophores for the Oaxacan coast, which represent new records of Beroe forskalii and Bolinopsis vitrea as well as the first record of Ocyropsis maculata in the tropical eastern Pacific. Descriptions of these three species, as well as a checklist of the ctenophores from the west coast of Mexico are provided.

Formation of the statolith in the ctenophore Mnemiopsis leidyi.

The aboral sensory organ (apical organ) of ctenophores contains a statocyst with a single large statolith. The statolith comprises living cells (lithocytes), each containing a large membrane-bound concretion. The statolith is supported on the distal ends of four compound motile mechanoresponsive cilia (balancers) which control the beat frequencies of the eight locomotory comb rows, and thereby the orientation of animals to gravity. In Mnemiopsis leidyi and Pleurobrachia pileus, lithocytes arise in the thickened epithelial floor of the apical organ on opposite sides along the tentacular plane. Lithocytes progressively differentiate and migrate toward the apical surface where they bud off next to the bases of the balancers. New lithocytes are transported up the balancers by ciliary surface motility to form the statolith (Noda, 2013). The statolith has a superellipsoidal shape due to the rectangular arrangement of the four balancers and the addition of new lithocytes to its ends via the balancers. The size of the statolith increases with animal size, starting at the highest rate of growth in younger stages and gradually decreasing in larger animals. The total number of developing lithocytes in the epithelial floor increases rapidly in smaller animals and reaches a plateau range in larger animals. Lithocytes are therefore produced continually throughout life for enlargement of the statolith and possibly for turnover and replacement of existing lithocytes. The dome cilia enclosing the statocyst were observed to propagate slow, low-ampitude waves distally. The dome cilia may act as an undulating screen to prevent foreign objects in the seawater from being transported non-specifically up the balancers to make a defective statolith.

Beroe gracilis (Ctenophora) from the Humboldt Current System: first occurrence of this species in the southern hemisphere.

Beroe gracilis Künne, 1939 is a small neritic ctenophore, previously recorded only from cold waters of the northern hemisphere. The present study provides the first record of the species in the southern hemisphere, found in the surface layer of the Humboldt Current System off the central Chilean coast (32°-36.5° S). A complete description of this material is provided.

Recurrent jellyfish blooms are a consequence of global oscillations.

A perceived recent increase in global jellyfish abundance has been portrayed as a symptom of degraded oceans. This perception is based primarily on a few case studies and anecdotal evidence, but a formal analysis of global temporal trends in jellyfish populations has been missing. Here, we analyze all available long-term datasets on changes in jellyfish abundance across multiple coastal stations, using linear and logistic mixed models and effect-size analysis to show that there is no robust evidence for a global increase in jellyfish. Although there has been a small linear increase in jellyfish since the 1970s, this trend was unsubstantiated by effect-size analysis that showed no difference in the proportion of increasing vs. decreasing jellyfish populations over all time periods examined. Rather, the strongest nonrandom trend indicated jellyfish populations undergo larger, worldwide oscillations with an approximate 20-y periodicity, including a rising phase during the 1990s that contributed to the perception of a global increase in jellyfish abundance. Sustained monitoring is required over the next decade to elucidate with statistical confidence whether the weak increasing linear trend in jellyfish after 1970 is an actual shift in the baseline or part of an oscillation. Irrespective of the nature of increase, given the potential damage posed by jellyfish blooms to fisheries, tourism, and other human industries, our findings foretell recurrent phases of rise and fall in jellyfish populations that society should be prepared to face.

Patterns of comb row development in young and adult stages of the ctenophores Mnemiopsis leidyi and Pleurobrachia pileus.

The development of comb rows in larval and adult Mnemiopsis leidyi and adult Pleurobrachia pileus is compared to regeneration of comb plates in these ctenophores. Late gastrula embryos and recently hatched cydippid larvae of Mnemiopsis have five comb plates in subsagittal rows and six comb plates in subtentacular rows. Subsagittal rows develop a new (sixth) comb plate and both types of rows add plates at similar rates until larvae reach the transition to the lobate form at ∼5 mm size. New plate formation then accelerates in subsagittal rows that later extend on the growing oral lobes to become twice the length of subtentacular rows. Interplate ciliated grooves (ICGs) develop in an aboral-oral direction along comb rows, but ICG formation itself proceeds from oral to aboral between plates. New comb plates in Mnemiopsis larvae are added at both aboral and oral ends of rows. At aboral ends, new plates arise as during regeneration: local widening of a ciliated groove followed by formation of a short split plate that grows longer and wider and joins into a common plate. At oral ends, new plates arise as a single tuft of cilia before an ICG appears. Adult Mnemiopsis continue to make new plates at both ends of rows. The frequency of new aboral plate formation varies in the eight rows of an animal and seems unrelated to body size. In Pleurobrachia that lack ICGs, new comb plates at aboral ends arise between the first and second plates as a single small nonsplit plate, located either on the row midline or off-axis toward the subtentacular plane. As the new (now second) plate grows larger, its distance from the first and third plates increases. Size of the new second plate varies within the eight rows of the same animal, indicating asynchronous formation of plates as in Mnemiopsis. New oral plates arise as in Mnemiopsis. The different modes of comb plate formation in Mnemiopsis versus Pleurobrachia are accounted for by differences in mesogleal firmness and mechanisms of ciliary coordination. In both cases, the body of a growing ctenophore is supplied with additional comb plates centripetally from opposite ends of the comb rows.

Mass occurrence of the ctenophore Bolinopsis vitrea (L. Agassiz, 1860) in the nearshore southern Adriatic Sea (Kotor Bay, Montenegro).

The ctenophore Bolinopsis vitrea has been rarely observed in the Mediterranean Sea. A bloom of B. vitrea is here reported for the first time in the southern Adriatic Sea in the spring and summer of 2009, together with its effect on the plankton of Kotor Bay. Ctenophores were found below 5 m depth only. Results of the investigation indicate that mass occurrence of B. vitrea could have a great impact on the Kotor Bay ecosystem. Their predation on copepods would reduce grazing pressure on phytoplankton, favouring an uncommon bloom of the latter. It is evident that B. vitrea are capable of altering rapidly the composition and biomass of coastal plankton communities when present in large masses. This first evidence of such events for this species may indicate changes in the functioning of marine ecosystems of the southern Adriatic.

Six major steps in animal evolution: are we derived sponge larvae?

A review of the old and new literature on animal morphology/embryology and molecular studies has led me to the following scenario for the early evolution of the metazoans. The metazoan ancestor, "choanoblastaea," was a pelagic sphere consisting of choanocytes. The evolution of multicellularity enabled division of labor between cells, and an "advanced choanoblastaea" consisted of choanocytes and nonfeeding cells. Polarity became established, and an adult, sessile stage developed. Choanocytes of the upper side became arranged in a groove with the cilia pumping water along the groove. Cells overarched the groove so that a choanocyte chamber was formed, establishing the body plan of an adult sponge; the pelagic larval stage was retained but became lecithotrophic. The sponges radiated into monophyletic Silicea, Calcarea, and Homoscleromorpha. Homoscleromorph larvae show cell layers resembling true, sealed epithelia. A homoscleromorph-like larva developed an archenteron, and the sealed epithelium made extracellular digestion possible in this isolated space. This larva became sexually mature, and the adult sponge-stage was abandoned in an extreme progenesis. This eumetazoan ancestor, "gastraea," corresponds to Haeckel's gastraea. Trichoplax represents this stage, but with the blastopore spread out so that the endoderm has become the underside of the creeping animal. Another lineage developed a nervous system; this "neurogastraea" is the ancestor of the Neuralia. Cnidarians have retained this organization, whereas the Triploblastica (Ctenophora+Bilateria), have developed the mesoderm. The bilaterians developed bilaterality in a primitive form in the Acoelomorpha and in an advanced form with tubular gut and long Hox cluster in the Eubilateria (Protostomia+Deuterostomia). It is indicated that the major evolutionary steps are the result of suites of existing genes becoming co-opted into new networks that specify new structures. The evolution of the eumetazoan ancestor from a progenetic homoscleromorph larva implies that we, as well as all the other eumetazoans, are derived sponge larvae.

Insights from diploblasts; the evolution of mesoderm and muscle.

The origin of both mesoderm and muscle are central questions in metazoan evolution. The majority of metazoan phyla are triploblasts, possessing three discrete germ layers. Attention has therefore been focused on two outgroups to triploblasts, Cnidaria and Ctenophora. Modern texts describe these taxa as diploblasts, lacking a mesodermal germ layer. However, some members of Medusozoa, one of two subphyla within Cnidaria, possess tissue independent of either the ectoderm or endoderm referred to as the entocodon. Furthermore, members of both Cnidaria and Ctenophora have been described as possessing striated muscle, a mesodermal derivative. While it is widely accepted that the ancestor of Eumetazoa was diploblastic, homology of the entocodon and mesoderm as well as striated muscle within Eumetazoa has been suggested. This implies a potential triploblastic ancestor of Eumetazoa possessing striated muscle. In the following review, I examine the evidence for homology of both muscle and mesoderm. Current data support a diploblastic ancestor of cnidarians, ctenophores, and triploblasts lacking striated muscle.