Feeding rates in sessile versus motile ciliates are hydrodynamically equivalent

  1. Jingyi Liu
  2. Yi Man
  3. John H Costello
  4. Eva Kanso  Is a corresponding author
  1. Department of Aerospace and Mechanical Engineering, University of Southern California, United States
  2. Mechanics and Engineering Science, Peking University, China
  3. Department of Biology, Providence College, United States
  4. Whitman Center, Marine Biological Laboratories, United States
  5. Department of Physics and Astronomy, University of Southern California, United States
8 figures, 4 tables and 1 additional file

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 (X,Y,Z).

Stokeslet and envelope models of sessile and motile ciliates.

(A) Stokeslet model where ciliary activity is represented by a Stokeslet force Fcilia is located at a distance (La)/a 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, L/a=2, a=1 and Fcilia is chosen to generate a swimming speed U=2/3 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 Q of a fluid volume passing through an annular disk of radius R/a=1.1 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 ΔQ=QmotileQsessile and Sherwood number ΔSh=ΔI/Idiffusion=ShmotileShsessile in the Stokeslet-sphere model for L/a=1.1 and L/a=2 and in the envelope model. In both metrics, the difference is less than 20%: ΔQ is less than 20% the advective flux πR2U and ΔI is less than 20% of the corresponding diffusive uptake Idiffusion=4πRDC. 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 D=1×109m2s1 and the solid symbols correspond to the diffusivity D=4×1010m2s1 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.

Appendix 1—figure 1
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 Vcylinder=πD2L/4,Vcone=πD2L/12, and Vspheroid=πW2L/6. Later on, we further simplify those shapes to an equal volume-based sphere with radius a, calculated as the formula shown in B for each geometry.

Appendix 1—figure 2
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 U is the swimming speed, while for sessile organisms, the characteristic speed U is the maximum flow speed reported near the organism. The characteristic length a is based on the volume-equivalent spherical radius. (A) Ratio of flow speed to length scale U/a in regular scale. (B) Ratio of flow speed to length scale U/a in logarithmic scale. (C) Advection strength aU in logarithmic scale.

Appendix 1—figure 3
Stokeslet model.

(A) Flow field around sessile and swimming spheres. Solution in non-dimensional form for a=1 and B=Fcilia/(8πμa)=1. (B) Fluid around a motile and sessile sphere with same point force strength at distance L=[2a,5a,10a], respectively. (C) Normalized clearance varies as annular encounter disk R for above same three point force distances.L=[2a,5a,10a]

Author response image 1

Tables

Table 1
Survey of size a and flow measurements U in sessile and swimming ciliates and sinking diatoms.

Size a is calculated using the volume-equivalent spherical radius. Corresponding ranges of Pe numbers are based on the diffusivity of oxygen, D=109m2s1, live bacteria, D=4×1010m2s1,, and dead bacteria D=2×1013m2s1.

Empirical measurementsPéclet number, Pe
microorganism sizecharacteristic speedoxygen diffusivitylive bacteria diffusivitydead bacteria diffusivity
Sessile ciliates15–6050–25001–802–210(5–400)×103
Swimming ciliates15–18050–32001–1608–390(17–800)×103
Sinking diatoms10–12040–2100.4–23
Appendix 1—table 1
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 r and angular variable μ=cosθ.

B.C. at the surface of the sphereB.C. at infinity
Sessile ciliated sphereuθ|r=a=B1μ2, ur|r=a=0u|r=0
Swimming ciliated sphereuθ|r=a=B1μ2, ur|r=a=0u|r=Uez
Sinking (non-ciliated) sphereu|r=a=0u|r=Uey
Fluid velocity field
Sessileur(r,μ)=(a3r3ar)Bμ,uθ(r,μ)=12(a3r3+ar)B1μ2
Swimmingur(r,μ)=(23+2a33r3)Bμ,uθ(r,μ)=(23+a33r3)B1μ2,
Sinkingur(r,μ)=(1+3a2ra32r3)Uμ,uθ(r,μ)=(13a4ra34r3)U1μ2
Fluid velocity field in lab frameFar-field signature
Sessilesame as aboveForce monopole (u1/r) (Stokeslet)
Swimmingur(r,μ)=2a33r3Bμ,uθ(r,μ)=a33r3B1μ2Potential dipole (u1/r3)
Sinkingur(r,μ)=(3a2ra32r3)Uμ,uθ(r,μ)=(3a4ra34r3)U1μ2,Force monopole (u1/r) (Stokeslet)
Appendix 1—table 2
Appendix 1—table 1 extension.
Fluid pressure fieldForces on sphere
Sessilep(r,μ)=pηar2BμF=4πηaBez
Swimmingp(r,μ)=pηar2BμF=4πηaBez, D=6πηaUez
Sinkingp(r,μ)=p+η3a2r2UμW=43πa3(δρ)gey, D=6πηaUey
Hydrodynamic powerSwimming speed
SessileP=8πaηB2U=0
SwimmingP=8πaη23B2U=23B
SinkingP=6πaηU2U=2ga2(δρ)9η
Appendix 1—table 3
Expressions for Sh as a function of Pe for sessile and swimming ciliated sphere model, compared to a sinking sphere.
Large Pe limitSmall Pe limit
Sherwood numberReferenceSherwood numberReference
SessileSh=23πPe12Present studySh=1+43720Pe2Present study
SwimmingSh=23πPe12Magar et al., 2003; Michelin and Lauga, 2011Sh=1+13PeMagar et al., 2003; Michelin and Lauga, 2011
SinkingSh=0.55Pe13Acrivos and Goddard, 1965; Acrivos and Taylor, 1962; Guasto et al., 2012Sh=1+13PeAcrivos and Taylor, 1962; Guasto et al., 2012

Additional files

Download links

A two-part list of links to download the article, or parts of the article, in various formats.

Downloads (link to download the article as PDF)

Open citations (links to open the citations from this article in various online reference manager services)

Cite this article (links to download the citations from this article in formats compatible with various reference manager tools)

  1. Jingyi Liu
  2. Yi Man
  3. John H Costello
  4. Eva Kanso
(2026)
Feeding rates in sessile versus motile ciliates are hydrodynamically equivalent
eLife 13:RP99003.
https://doi.org/10.7554/eLife.99003.3