Microbial filter feeders are an important group of grazers, significant to the microbial loop, aquatic food webs, and biogeochemical cycling. Our understanding of microbial filter feeding is poor, and, importantly, it is unknown what force microbial filter feeders must generate to process adequate amounts of water. Also, the trade-off in the filter spacing remains unexplored, despite its simple formulation: A filter too coarse will allow suitably sized prey to pass unintercepted, whereas a filter too fine will cause strong flow resistance. We quantify the feeding flow of the filter-feeding choanoflagellate Diaphanoeca grandis using particle tracking, and demonstrate that the current understanding of microbial filter feeding is inconsistent with computational fluid dynamics (CFD) and analytical estimates. Both approaches underestimate observed filtration rates by more than an order of magnitude; the beating flagellum is simply unable to draw enough water through the fine filter. We find similar discrepancies for other choanoflagellate species, highlighting an apparent paradox. Our observations motivate us to suggest a radically different filtration mechanism that requires a flagellar vane (sheet), something notoriously difficult to visualize but sporadically observed in the related choanocytes (sponges). A CFD model with a flagellar vane correctly predicts the filtration rate of D. grandis, and using a simple model we can account for the filtration rates of other microbial filter feeders. We finally predict how optimum filter mesh size increases with cell size in microbial filter feeders, a prediction that accords very well with observations. We expect our results to be of significance for small-scale biophysics and trait-based ecological modeling. protozoans | choanoflagellates | filter feeding | microswimmers | computational fluid dynamics H eterotrophic microorganisms in the oceans inhabit a dilute environment and they need efficient feeding mechanisms to acquire enough food to sustain growth (1, 2). At the microscale the Reynolds number is low and viscous forces govern hydrodynamical interactions. This implies extensive, long-range flow disturbances around moving particles and microswimmers, impeding cell-cell contact and prey capture (3, 4). However, to encounter enough food, purely heterotrophic plankton that rely solely on prey capture typically need to clear a volume of water for prey corresponding to 1 million times their own body volume per day (4). Thus, heterotrophic microbes face a difficult challenge, and the prevailing viscous forces must strongly influence prey capture and shape the various feeding modes through evolution.Many unicellular flagellates as well as colonial sponges and metazoans, e.g., tunicates, use filter feeding to catch bacteriasized prey (1, 5-7). They establish a feeding current, from which prey particles are sieved using filter structures. Such filter feeders benefit from having filters with small mesh size that allow the organisms to capture small prey (5, 8). However, filter ...
Leuconoid sponges are filter-feeders with a complex system of branching inhalant and exhalant canals leading to and from the close-packed choanocyte chambers. Each of these choanocyte chambers holds many choanocytes that act as pumping units delivering the relatively high pressure rise needed to overcome the system pressure losses in canals and constrictions. Here, we test the hypothesis that, in order to deliver the high pressures observed, each choanocyte operates as a leaky, positive displacement-type pump owing to the interaction between its beating flagellar vane and the collar, open at the base for inflow but sealed above. The leaking backflow is caused by small gaps between the vaned flagellum and the collar. The choanocyte pumps act in parallel, each delivering the same high pressure, because low-pressure and high-pressure zones in the choanocyte chamber are separated by a seal (secondary reticulum). A simple analytical model is derived for the pump characteristic, and by imposing an estimated system characteristic we obtain the back-pressure characteristic that shows good agreement with available experimental data. Computational fluid dynamics is used to verify a simple model for the dependence of leak flow through gaps in a conceptual collar–vane–flagellum system and then applied to models of a choanocyte tailored to the parameters of the freshwater demosponge Spongilla lacustris to study its flows in detail. It is found that both the impermeable glycocalyx mesh covering the upper part of the collar and the secondary reticulum are indispensable features for the choanocyte pump to deliver the observed high pressures. Finally, the mechanical pump power expended by the beating flagellum is compared with the useful (reversible) pumping power received by the water flow to arrive at a typical mechanical pump efficiency of about 70%.
Sponges are suspension feeders that filter vast amounts of water. Pumping is carried out by flagellated chambers that are connected to an inhalant and exhalant canal system. In 'leucon' sponges with relatively high-pressure resistance due to a complex and narrow canal system, pumping and filtering are only possible owing to the presence of a gasket-like structure (forming a canopy above the collar filters). Here we combine numerical and experimental work, and demonstrate how sponges that lack such sealing elements are able to efficiently pump and force the flagella driven flow through their collar filter, thanks to the formation of a 'hydrodynamic gasket' above the collar. Our findings link the architecture of flagellated chambers to that of the canal system, and lend support to the current view that the sponge aquiferous system evolved from an open-type filtration system, and that the first metazoans were filter feeders.
Wave propagation of carbon nanotubes embedded in an elastic medium J. Appl. Phys. 97, 044307 (2005); 10.1063/1.1849823Simulation study of carbon nanotube field emission display with under-gate and planar-gate structures Carbon nanotube webs exhibit interesting properties when used as thermo-acoustic projectors. This work studies thermo-acoustic effect of these sound sources both in near and far field regions. Based on two alternative forms of the energy equation, we have developed a straightforward formula for calculation of pressure field, which is consistent with experimental data in far field. Also we have solved full 3-D governing equations using numerical methods. Our three-dimensional simulation and experimental data show pressure waves are highly affected by dimensions of sound sources in near field due to interference effects. However, generation of sound waves in far field is independent of projectors area surface. Energy analysis for free standing Thermo-Acoustic (TA) sound sources show that aerogel TA sound sources like CNT based projectors could act more efficiently compared to the other sources in delivering more than 75% of alternative input energy to the medium gas up to a frequency of 1 MHz. V C 2015 AIP Publishing LLC.
Choanoflagellates are unicellular eukaryotes that are ubiquitous in aquatic habitats. They have a single flagellum that creates a flow toward a collar filter composed of filter strands that extend from the cell. In one common group, the loricate choanoflagellates, the cell is suspended in an elaborate basket-like structure, the lorica, the function of which remains unknown. Here, we use Computational Fluid Dynamics to explore the possible hydrodynamic function of the lorica. We use the choanoflagellate Diaphaoneca grandis as a model organism. It has been hypothesized that the function of the lorica is to prevent refiltration (flow recirculation) and to increase the drag and, hence, increase the feeding rate and reduce the swimming speed. We find no support for these hypotheses. On the contrary, motile prey are encountered at a much lower rate by the loricate organism. The presence of the lorica does not affect the average swimming speed, but it suppresses the lateral motion and rotation of the cell. Without the lorica, the cell jiggles from side to side while swimming. The unsteady flow generated by the beating flagellum causes reversed flow through the collar filter that may wash away captured prey while it is being transported to the cell body for engulfment. The lorica substantially decreases such flow, hence it potentially increases the capture efficiency. This may be the main adaptive value of the lorica.
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