Abstract
The clams, mussels, and tubeworms that dominate deep-sea chemosynthetic communities obtain most of their nutrition through intracellular symbiotic g-Proteobacteria that oxidize reduced compounds. The modes of symbiont transmission employed by various taxa have profound consequences for genetic, demographic, and evolutionary processes affecting the symbionts and their hosts. Vesicomyid clams transmit endosymbionts vertically via their eggs, a process that leads to symbiont clonality and accelerated rates of evolution. Vertical transmission provides the host with symbiont assurance because dispersing larvae carry the bacteria as they colonize new habitats. The symbionts and their host clams exhibit cospeciation. Vertical transmission for at least 45 million years has contributed to significant genome reduction, as the symbionts have lost almost half their DNA and many of the genes that were required for living in the ambient environment. In contrast, the horizontally transmitted symbionts associated with siboglinid tubeworms do not exhibit genome reduction. Tubeworm larvae are newly infected in each generation when they settle on appropriate substrates. Infection by local bacterial strains is hypothesized to provide the worms with locally optimal symbionts. Symbiont diversity is structured geographically and by habitat type (vent vs seep) and does not parallel host evolution. Less is known about the endosymbionts associated with various species of bathymodiolin mussels. Acquisition of local symbionts occurs in these mussels, but a vertical component of transmission might also exist. Symbiont diversity is structured geographically and not according to host species. The benefits of various symbiont transmission modes also carry associated risks that range from pure enslavement and genomic erosion under strictly vertical transmission to the possible evolution of bacterial strains that cheat the host when mixed symbiont genotypes infect a single host under horizontal transmission. The prevalence of horizontal transmission systems in chemosynthetic environments suggests that the symbionts must have escape strategies that allow them to re-inoculate the ambient environment and contribute to their overall fitness.
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Notes
- 1.
The quotes denote the dubious assignment of this genus name to these species (Jones and Vrijenhoek 2006).
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Acknowledgements
I wish to thank the Frank Stewart, Shana Goffredi, Monika Bright, Julie Robidart, Steffen Kiel, Julio Harvey, Shannon Johnson and an anonymous reviewer for providing information and criticisms that improved the scope of this manuscript. Funding was provided by grants from the David and Lucile Packard Foundation to the Monterey Bay Research Institute and the National Science Foundation (OCE 0241613).
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Vrijenhoek, R.C. (2010). Genetics and Evolution of Deep-Sea Chemosynthetic Bacteria and Their Invertebrate Hosts. In: Kiel, S. (eds) The Vent and Seep Biota. Topics in Geobiology, vol 33. Springer, Dordrecht. https://doi.org/10.1007/978-90-481-9572-5_2
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DOI: https://doi.org/10.1007/978-90-481-9572-5_2
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