Bivalve specimens from legacy frozen tissue collections, and others freshly obtained, were surveyed for the presence of the Steamer long terminal repeat (LTR)-retrotransposon associated with disseminated hemic neoplasia of the soft-shelled clam Mya areneria. Of 22 species investigated using primers for the pol region, only Atlantic M. arenaria, Atlantic and North Sea razor clams Ensis directus, and Baltic clams Macoma balthica from the North Sea were found to possess copies of Steamer in their genomes. Notably, close relatives like Mya truncata and Siliqua patula did not exhibit evidence of Steamer. Amplified Steamer sequences were uniformly identical in all M. areneria specimens, and were highly variable across specimens of E. directus. Variation in the latter included nucleotide polymorphisms among and within individuals as well as length variation in 2 specimens corresponding to the deletion of a predicted stable hairpin structure. Results implicate Atlantic razor clams as the proximal source for horizontal transmission of Steamer among ecologically similar yet markedly distantly related bivalves. The consequences of cross-species transmission of the Steamer retrotransposon are unknown, and the finding of Steamer in 3 bivalve species suggests that further spread is possible.
KEY WORDS: Disseminated neoplasia · Hemic neoplasia · Retrotransposon · Steamer · Bivalvia · CancerResale or republication not permitted without written consent of the publisher Dis Aquat Org 124: [165][166][167][168] 2017 pression of the Steamer retrotransposon. Steamer is a long terminal repeat (LTR)-retrotrans poson in the Mag family of Ty3/Gypsy elements, complete with a single open reading frame encoding gag and pol genes. Retrotransposons are present in genomic DNA and replicate through reverse transcription of an mRNA transcript into DNA, followed by integration into a new location within the genomic DNA of a cell. As they are notably lacking a retroviral env gene, they are not expected to be able to transfer from cell to cell, as retroviruses do. Similar retrotransposons have been identified in the genomes of 3 other bivalve species known to have transmissible neoplasias, one of which seems to cross the species boundary (Metzger et al. 2016).To date, the Steamer retrotransposon itself is un ique to M. arenaria. Here we examined legacy bio diversity collections and freshly acquired bivalve tissues for the presence of a Steamer signature in their genomes in a manner that might (1) better elucidate whether disseminated neoplasias are ecologically or phylogenetically determined and (2) shed light on the potential for horizontal spread and genomic modification by Steamerlike retrotransposons in the marine environment.
MATERIALS AND METHODSBivalve tissue samples were obtained from the American Museum of Natural History's Ambrose Monell Cryo Collection (AMCC), and included 22 species from the North Atlantic (see Table S1 in the Supplement at www. int-res. com/ articles/ suppl/ d124 p165 _ supp. pdf). Fresh specimens of steam...