Foraging mechanisms in excavate flagellates shed light on the functional ecology of early eukaryotes.

The phagotrophic flagellates described as "typical excavates" have been hypothesized to be morphologically similar to the Last Eukaryotic Common Ancestor and understanding the functional ecology of excavates may therefore help shed light on the ecology of these early eukaryotes. Typical excavates are characterized by a posterior flagellum equipped with a vane that beats in a ventral groove. Here, we combined flow visualization and observations of prey capture in representatives of the three clades of excavates with computational fluid dynamic modeling, to understand the functional significance of this cell architecture. We record substantial differences amongst species in the orientation of the vane and the beat plane of the posterior flagellum. Clearance rate magnitudes estimated from flow visualization and modeling are both like that of other similarly sized flagellates. The interaction between a vaned flagellum beating in a confinement is modeled to produce a very efficient feeding current at low energy costs, irrespective of the beat plane and vane orientation and of all other morphological variations. Given this predicted uniformity of function, we suggest that the foraging systems of typical excavates studied here may be good proxies to understand those potentially used by our distant ancestors more than 1 billion years ago.

Incorporating recirculation effects into metrics of feeding performance for current-feeding zooplankton.

The feeding performance of zooplankton influences their evolution and can explain their behaviour. A commonly used metric for feeding performance is the volume of fluid that flows through a filtering surface and is scanned for food. Here, we show that such a metric may give incorrect results for organisms that produce recirculatory flows, so that fluid flowing through the filter may have been already filtered of food. In a numerical model, we construct a feeding metric that correctly accounts for recirculation in a sessile model organism inspired by our experimental observations of Vorticella and its flow field. Our metric tracks the history of current-borne particles to determine if they have already been filtered by the filtering surface. Examining the pathlines of food particles reveals that the capture of fresh particles preferentially involves the tips of cilia, which we corroborate in observations of feeding Vorticella. We compare the amount of fresh nutrient particles carried to the organism with other metrics of feeding, and show that metrics that do not take into account the history of particles cannot correctly compute the volume of freshly scanned fluid.

Freshwater snail feeding: lubrication-based particle collection on the water surface.

The means by which aquatic animals such as freshwater snails collect food particles distributed on the water surface are of great interest for understanding life at the air-water interface. The apple snail Pomacea canaliculata stabilizes itself just below the air-water interface and manipulates its foot such that it forms a cone-shaped funnel into which an inhalant current is generated, thereby drawing food particles into the funnel to be ingested. We measured the velocity of this feeding current and tracked the trajectories of food particles around and on the snail. Our experiments indicated that the particles were collected via the free surface flow generated by the snail's undulating foot. The findings were interpreted using a simple model based on lubrication theory, which considered several plausible mechanisms depending on the relative importance of hydrostatic pressure, capillary action and rhythmic surface undulation.

Sulfide assimilation by ectosymbionts of the sessile ciliate, Zoothamnium niveum.

We investigated the constraints on sulfide uptake by bacterial ectosymbionts on the marine peritrich ciliate Zoothamnium niveum by a combination of experimental and numerical methods. Protists with symbionts were collected on large blocks of mangrove-peat. The blocks were placed in a flow cell with flow adjusted to in situ velocity. The water motion around the colonies was then characterized by particle tracking velocimetry. This shows that the feather-shaped colony of Z. niveum generates a unidirectional flow of seawater through the colony with no recirculation. The source of the feeding current was the free-flowing water although the size of the colonies suggests that they live partly submerged in the diffusive boundary layer. We showed that the filtered volume allows Z. niveum to assimilate sufficient sulfide to sustain the symbiosis at a few micromoles per liter in ambient concentration. Numerical modeling shows that sulfide oxidizing bacteria on the surfaces of Z. niveum can sustain 100-times higher sulfide uptake than bacteria on flat surfaces, such as microbial mats. The study demonstrates that the filter feeding zooids of Z. niveum are preadapted to be prime habitats for sulfide oxidizing bacteria due to Z. niveum's habitat preference and due to the feeding current. Z. niveum is capable of exploiting low concentrations of sulfide in near norm-oxic seawater. This links its otherwise dissimilar habitats and makes it functionally similar to invertebrates with thiotrophic symbionts in filtering organs.