We describe development of the hoplonemertean Paranemertes peregrina from fertilization to juvenile, using light, confocal, and electron microscopy. We discovered that the uniformly ciliated lecithotrophic larva of this species has a transitory epidermis, which is gradually replaced by the definitive epidermis during the course of planktonic development. The approximately 90 large multiciliated cleavage-arrested cells of the transitory larval epidermis become separated from each other by intercalating cells of the definitive epidermis, then gradually diminish in size and disappear more or less simultaneously. Rudiments of all major adult structures-the gut, proboscis, cerebral ganglia, lateral nerve cords, and cerebral organs-are already present in 4-day-old larvae. Replacement of the epidermis is the only overt metamorphic transformation of larval tissue; larval structures otherwise prefigure the juvenile body, which is complete in about 10 days at 7-10 degrees C. Our findings on development of digestive system, nervous system, and proboscis differ in several ways from previous descriptions of hoplonemertean development. We report development with transitory epidermis in two other species, review evidence from the literature, and suggest that this developmental type is the rule for hoplonemerteans. The hoplonemertean planuliform larva is fundamentally different both from the pilidium larva of the sister group to the Hoplonemertea, the Pilidiophora, and from the hidden trochophore of palaeonemerteans. We discuss the possible function and homology of the larval epidermis in development of other nemerteans and spiralians in general.
Background: The new animal phylogeny established several taxa which were not identified by morphological analyses, most prominently the Ecdysozoa (arthropods, roundworms, priapulids and others) and Lophotrochozoa (molluscs, annelids, brachiopods and others). Lophotrochozoan interrelationships are under discussion, e.g. regarding the position of Nemertea (ribbon worms), which were discussed to be sister group to e.g. Mollusca, Brachiozoa or Platyhelminthes. Mitochondrial genomes contributed well with sequence data and gene order characters to the deep metazoan phylogeny debate.
Nemerteans (ribbon worms) employ toxins to subdue their prey, but research thus far has focused on the small-molecule components of mucus secretions and few protein toxins have been characterized. We carried out a preliminary proteotranscriptomic analysis of putative toxins produced by the hoplonemertean Amphiporus lactifloreus (Hoplonemertea, Amphiporidae). No variants were found of known nemertean-specific toxin proteins (neurotoxins, cytotoxins, parbolysins or nemertides) but several toxin-like transcripts were discovered, expressed strongly in the proboscis, including putative metalloproteinases and sequences resembling sea anemone actitoxins, crown-of-thorn sea star plancitoxins, and multiple classes of inhibitor cystine knot/knottin family proteins. Some of these products were also directly identified in the mucus proteome, supporting their preliminary identification as secreted toxin components. Two new nemertean-typical toxin candidates could be described and were named U-nemertotoxin-1 and U-nemertotoxin-2. Our findings provide insight into the largely overlooked venom system of nemerteans and support a hypothesis in which the nemertean proboscis evolved in several steps from a flesh-melting organ in scavenging nemerteans to a flesh-melting and toxin-secreting venom apparatus in hunting hoplonemerteans.
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