Bacterial symbiosis in Syssitomya pourtalesiana Oliver, 2012 (Galeommatoidea: Montacutidae), a bivalve commensal with the deep-sea echinoid Pourtalesia.

For the first time, bacterial symbiosis is recognized in the bivalve family Montacutidae of the superfamily Galeommatoidea. The ctenidial filaments of Syssitomya pourtalesiana Oliver, 2012 are extended abfrontally and a dense layer of bacteriocyte cells cover the entire surface behind a narrow ciliated frontal zone. The bacteria are extracellular and held within a matrix of epithelial extensions and microvilli. There is no cuticular layer (glycocalyx) covering the bacteria as in many thyasirid symbioses. The bacteriocytes hold more than one morphotype of bacteria, but bacilli, 1–3 μm in length, dominate. Scanning electron microscopy observations show a surface mat of filamentous bacteria over the extreme abfrontal surfaces. Filter feeding was confirmed by the presence of food particles in the stomach and the bivalve is presumed to be mixotrophic. Syssitomya is commensal and lives attached to the anal spines of the deep-sea echinoid Pourtalesia. In this position, echinoid feeding currents and echinoid faecal material may supply the bacteria with a variety of nutrient materials including dissolved organic matter.

Implications of streamlining theory for microbial ecology

Whether a small cell, a small genome or a minimal set of chemical reactions with self-replicating properties, simplicity is beguiling. As Leonardo da Vinci reportedly said, 'simplicity is the ultimate sophistication'. Two diverging views of simplicity have emerged in accounts of symbiotic and commensal bacteria and cosmopolitan free-living bacteria with small genomes. The small genomes of obligate insect endosymbionts have been attributed to genetic drift caused by small effective population sizes (Ne). In contrast, streamlining theory attributes small cells and genomes to selection for efficient use of nutrients in populations where Ne is large and nutrients limit growth. Regardless of the cause of genome reduction, lost coding potential eventually dictates loss of function. Consequences of reductive evolution in streamlined organisms include atypical patterns of prototrophy and the absence of common regulatory systems, which have been linked to difficulty in culturing these cells. Recent evidence from metagenomics suggests that streamlining is commonplace, may broadly explain the phenomenon of the uncultured microbial majority, and might also explain the highly interdependent (connected) behavior of many microbial ecosystems. Streamlining theory is belied by the observation that many successful bacteria are large cells with complex genomes. To fully appreciate streamlining, we must look to the life histories and adaptive strategies of cells, which impose minimum requirements for complexity that vary with niche.