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Disseminated neoplasia (DN) is a form of cancer in bivalve molluscs that can be transmissible between individuals and in some cases across species. Neoplastic cells are highly proliferative, and infection is usually lethal. Commercially valuable bivalve species (mussels, cockles, softshell clams, and oysters) are affected by outbreaks of DN, making disease diagnosis and mitigation an important issue in ecological restoration efforts and aquaculture. Basket cockles (Clinocardium nuttallii) are native to the North American Pacific coast from California to Alaska. Recent concern from some Coast Salish Tribes regarding an observed long-term decline in basket cockle populations in Puget Sound, WA has increased interest in monitoring efforts and subsequent collection for aquarium-reared broodstock. Disseminated neoplasia was detected in Puget Sound basket cockle populations, delaying aquaculture efforts so that potential broodstock could be assessed for the presence of DN. This study details a minimally invasive, inexpensive, nonlethal method for high-throughput screening for DN in adult basket cockles. The hemolymph smear screening method to diagnose DN in C. nuttallii can be applied at field sites at low financial cost. Results of the hemolymph smear technique were validated against whole tissue histology, the standard method for DN diagnosis. Due to the similar cellular morphologies of DN in different bivalve species, it is proposed that hemolymph histology can likely be applied for diagnosing DN in other bivalves. 

Abstract Transmissible cancers are infectious parasitic clones that metastasize to new hosts, living past the death of the founder animal in which the cancer initiated. We investigated the evolutionary history of a cancer lineage that has spread though the soft-shell clam (Mya arenaria) population by assembling a chromosome-scale soft-shell clam reference genome and characterizing somatic mutations in transmissible cancer. We observe high mutation density, widespread copy-number gain, structural rearrangement, loss of heterozygosity, variable telomere lengths, mitochondrial genome expansion and transposable element activity, all indicative of an unstable cancer genome. We also discover a previously unreported mutational signature associated with overexpression of an error-prone polymerase and use this to estimate the lineage to be >200 years old. Our study reveals the ability for an invertebrate cancer lineage to survive for centuries while its genome continues to structurally mutate, likely contributing to the evolution of this lineage as a parasitic cancer. 

Many pathogens can cause cancer, but cancer itself does not normally act as an infectious agent. However, transmissible cancers have been found in a few cases in nature: in Tasmanian devils, dogs, and several bivalve species. The transmissible cancers in dogs and devils are known to spread through direct physical contact, but the exact route of transmission of bivalve transmissible neoplasia (BTN) has not yet been confirmed. It has been hypothesized that cancer cells from bivalves could be released by diseased animals and spread through the water column to infect/engraft into other animals. To test the feasibility of this proposed mechanism of transmission, we tested the ability of BTN cells from the soft-shell clam (Mya arenaria BTN, or MarBTN) to survive in artificial seawater. We found that MarBTN cells are highly sensitive to salinity, with acute toxicity at salinity levels lower than those found in the native marine environment. BTN cells also survive longer at lower temperatures, with 50% of cells surviving greater than 12 days in seawater at 10 °C, and more than 19 days at 4 °C. With one clam donor, living cells were observed for more than eight weeks at 4 °C. We also used qPCR of environmental DNA (eDNA) to detect the presence of MarBTN-specific DNA in the environment. We observed release of MarBTN-specific DNA into the water of laboratory aquaria containing highly MarBTN-diseased clams, and we detected MarBTN-specific DNA in seawater samples collected from MarBTN-endemic areas in Maine, although the copy numbers detected in environmental samples were much lower than those found in aquaria. Overall, these data show that MarBTN cells can survive well in seawater, and they are released into the water by diseased animals. These findings support the hypothesis that BTN is spread from animal-to-animal by free cells through seawater.