Size dependence of composition, photosynthesis and growth in the colony-forming freshwater ciliate, Ophrydium versatile

Abstract

SUMMARY 1. Ophrydium versatile is a symbiotic ciliate which forms gelatinous colonies up to several centimetres in diameter in transparent temperate lakes. The ciliates are evenly spaced at the colony surface and constitute a small proportion of the surface area (7%) and volume (3.1%) of the colony, but a large proportion of organic carbon (74%) and nitrogen content (82%) (exemplified for 1 cm3 colonies). The majority of the colony volume is formed by the jelly. The biomass proportion of ciliates scales inversely with colony size, following the decline of surface area to colony volume. The largest colonies found in Danish lakes in early summer contain almost 1 million ciliates, and assuming they derive from a single ciliate undergoing exponential division, they need twenty generations and, presumably, almost a year to reach maximum size. 2. The ciliates contain numerous symbiotic zoochlorellae that constitute about 10% of ciliate volume and more than half of the carbon content. Zoochlorellae dominate oxygen metabolism of the assemblage, resulting in low light compensation points, a large diel photosynthetic surplus, and a marked dependence on light for sustained growth and ciliate metabolism. Estimated gross photosynthesis (7ng C ciliate−1 day−1) of Ophnrydium from shallow, clear waters in June greatly exceeded the estimated carbon contained in filtered bacteria and small algae (1.9ng C cilicate−1 day−1). Nitrogen and phosphorus content of the prey, however, may provide the main nutrient source consistent with the correspondence between mass-specific rates of nutrient uptake and measured relative growth rates (average 0.067 day−1, generation time 10 days). 3. The large Ophrydium colonies require increased allocation of photosynthetic carbohydrates with increasing colony size to maintain the jelly. The large colonies tend to become gas-filled, floating, mechanically destroyed and their ciliate inhabitants abandon them as swarmers. Colony formation, however, should offer protection against predators which may be more important for the natural abundance than the costs of growing in a colony.