Feeding rates in sessile versus motile ciliates are hydrodynamically equivalent
Figures
Phylogenetic tree.
(A) Phylogenetic tree showing microorganisms known to feature cilia that generate feeding currents in either sessile (blue) or free swimming (purple) states. The class of diatoms – non-motile cells that sink when experiencing nutrient limitation – is shown for comparison. (B) Flow fields around a sessile ciliate, swimming ciliate, and sinking diatom, in lab and body frame of references. Streamlines are shown in blue in the lab frame .
Stokeslet and envelope models of sessile and motile ciliates.
(A) Stokeslet model where ciliary activity is represented by a Stokeslet force is located at a distance outside the spherical cell surface with no-slip surface velocity. (B) Envelope model where cilia activity is distributed over the entire cell surface with slip surface velocity. (C, D) Fluid streamlines (white) and nutrient concentration fields (colormap) in the sessile and swimming cases. Here, , and is chosen to generate a swimming speed in the motile case to ensure consistency with the envelope model. (E, F) Nutrient uptake in sessile and motile Stokeslet-sphere model based on calculation of clearance rate of a fluid volume passing through an annular disk of radius and Sherwood number Sh. In the latter, Pe is 100. (G) Nutrient uptake in sessile and motile envelope model based on calculation of Sherwood number Sh as a function of Pe. (H) Difference in clearance rate and Sherwood number in the Stokeslet-sphere model for and and in the envelope model. In both metrics, the difference is less than 20%: is less than 20% the advective flux and is less than 20% of the corresponding diffusive uptake . The shaded gray area denotes when the sessile strategy is advantageous.
Sherwood number versus Péclet number for the sinking (green) diatom and the swimming (purple) and sessile (blue) ciliates based on the envelope model.
(A) Shifted Sherwood number (Sh - 1) versus Péclet number in the logarithmic scale for a range of Pe from 0 to 1000. Pe numbers associated with experimental observations of diatoms (square), swimming ciliates (triangle), and sessile ciliates (circle) are superimposed. Corresponding Sh numbers are calculated based on the mathematical model. Empty symbols are for oxygen diffusivity and the solid symbols correspond to the diffusivity of live bacteria (Berg, 2018). (B–C) Asymptotic analysis (dashed lines) of Sherwood number in the large Péclet limit (B) and small Péclet limit (C).
Robustness to variations in cilia coverage and absorption fraction.
We considered a 50% cilia coverage and 50% absorption fraction located at back, middle, and front of the (A) sessile and (B) motile sphere. Concentration fields and Sherwood numbers with 100% cilia coverage and absorption area are shown in the top right corner. In all other cases, the Sh number is reported as a percentage of the full coverage/absorption case.
Geometry of ciliates and diatoms.
(A) Representative morphologies of surveyed species of sessile ciliates (blue), swimming ciliates (purple), and sinking diatoms (green). (B) Simplified shapes of above organisms. The volumes of those shapes can be obtained as ,, and . Later on, we further simplify those shapes to an equal volume-based sphere with radius , calculated as the formula shown in B for each geometry.
Raw data of character flow speed and size for sessile (blue) and swimming (purple) ciliates, and sinking diatoms (green).
For motile organisms, the characteristic flow speed is the swimming speed, while for sessile organisms, the characteristic speed is the maximum flow speed reported near the organism. The characteristic length is based on the volume-equivalent spherical radius. (A) Ratio of flow speed to length scale in regular scale. (B) Ratio of flow speed to length scale in logarithmic scale. (C) Advection strength in logarithmic scale.
Stokeslet model.
(A) Flow field around sessile and swimming spheres. Solution in non-dimensional form for and . (B) Fluid around a motile and sessile sphere with same point force strength at distance , respectively. (C) Normalized clearance varies as annular encounter disk for above same three point force distances.
Tables
Survey of size and flow measurements in sessile and swimming ciliates and sinking diatoms.
Size is calculated using the volume-equivalent spherical radius. Corresponding ranges of Pe numbers are based on the diffusivity of oxygen, , live bacteria, ,, and dead bacteria .
| Empirical measurements | Péclet number, Pe | ||||
|---|---|---|---|---|---|
| microorganism size | characteristic speed | oxygen diffusivity | live bacteria diffusivity | dead bacteria diffusivity | |
| Sessile ciliates | 15–60 | 50–2500 | 1–80 | 2–210 | (5–400)×103 |
| Swimming ciliates | 15–180 | 50–3200 | 1–160 | 8–390 | (17–800)×103 |
| Sinking diatoms | 10–120 | 40–210 | 0.4–23 | – | – |
Envelope model subject to treadmill slip velocity.
Mathematical expressions of boundary conditions, fluid velocity field, pressure field, forces acting on the sphere, hydrodynamic power, and speed for each representative case: sessile ciliated sphere, freely swimming ciliated sphere, and sinking (non-ciliated) sphere. All quantities are given in dimensional form in terms of the radial distance and angular variable .
| B.C. at the surface of the sphere | B.C. at infinity | |
|---|---|---|
| Sessile ciliated sphere | , | |
| Swimming ciliated sphere | , | |
| Sinking (non-ciliated) sphere | ||
| Fluid velocity field | ||
| Sessile | ||
| Swimming | ||
| Sinking | ||
| Fluid velocity field in lab frame | Far-field signature | |
| Sessile | same as above | Force monopole () (Stokeslet) |
| Swimming | Potential dipole () | |
| Sinking | , | Force monopole () (Stokeslet) |
Appendix 1—table 1 extension.
| Fluid pressure field | Forces on sphere | |
|---|---|---|
| Sessile | ||
| Swimming | , | |
| Sinking | , | |
| Hydrodynamic power | Swimming speed | |
| Sessile | ||
| Swimming | ||
| Sinking |
Expressions for Sh as a function of Pe for sessile and swimming ciliated sphere model, compared to a sinking sphere.
| Large Pe limit | Small Pe limit | |||
|---|---|---|---|---|
| Sherwood number | Reference | Sherwood number | Reference | |
| Sessile | Present study | Present study | ||
| Swimming | Magar et al., 2003; Michelin and Lauga, 2011 | Magar et al., 2003; Michelin and Lauga, 2011 | ||
| Sinking | Acrivos and Goddard, 1965; Acrivos and Taylor, 1962; Guasto et al., 2012 | Acrivos and Taylor, 1962; Guasto et al., 2012 | ||