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A swimming bacterium in a two-fluid model of a polymer solution

Published online by Cambridge University Press:  05 December 2024

Sabarish V. Narayanan
Affiliation:
Robert Frederick Smith School of Chemical and Biomolecular Engineering, Cornell University, Ithaca, NY 14853, USA
Donald L. Koch
Affiliation:
Robert Frederick Smith School of Chemical and Biomolecular Engineering, Cornell University, Ithaca, NY 14853, USA
Sarah Hormozi*
Affiliation:
Robert Frederick Smith School of Chemical and Biomolecular Engineering, Cornell University, Ithaca, NY 14853, USA
*
Email address for correspondence: hormozi@cornell.edu

Abstract

We analyse the motion of a flagellated bacterium in a two-fluid medium using slender body theory. The two-fluid model is useful for describing a body moving through a complex fluid with a microstructure whose length scale is comparable to the characteristic scale of the body. This is true for bacterial motion in biological fluids (entangled polymer solutions), where the entanglement results in a porous microstructure with typical pore diameters comparable to or larger than the flagellar bundle diameter, but smaller than the diameter of the bacterial head. Thus, the polymer and solvent satisfy different boundary conditions on the flagellar bundle and move with different velocities close to it. This gives rise to a screening length $L_B$ within which the fluids exchange momentum and the relative velocity between the two fluids decays. In this work, both the solvent and polymer of the two-fluid medium are modelled as Newtonian fluids with different viscosities $\mu _s$ and $\mu _p$ (viscosity ratio $\lambda = \mu _p/\mu _s$), thereby capturing the effects solely introduced by the microstructure of the complex fluid. From our calculations, we observe an increased drag anisotropy for a rigid, slender flagellar bundle moving through this two-fluid medium, resulting in an enhanced swimming velocity of the organism. The results are sensitive to the interaction between the bundle and the polymer, and we discuss two physical scenarios corresponding to two types of interaction. Our model provides an explanation for the experimentally observed enhancement of swimming velocity of bacteria in entangled polymer solutions and motivates further experimental investigations.

Information

Type
JFM Papers
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
Copyright
© The Author(s), 2024. Published by Cambridge University Press.
Figure 0

Figure 1. Plots of (a) drag force normalised by $F_N = 6 {\rm \pi} U \mu _s a$ on a sphere of radius $a$ translating with velocity $U$ and (b) torque normalised by $T_N = 8 {\rm \pi} \mu _s a^3 \omega$ on a sphere rotating with angular velocity $\omega$ in a two-fluid medium with a slipping polymer, as a function of $L_B/a$.

Figure 1

Figure 2. Plots of (a) drag force normalised by $F_N = 6 {\rm \pi} U \mu _s a$ on a sphere of radius $a$ translating with velocity $U$ and (b) torque normalised by $T_N = 8 {\rm \pi} \mu _s a^3 \omega$ on a sphere rotating with angular velocity $\omega$ in a two-fluid medium with no polymer–sphere interaction, as a function of $L_B/a$.

Figure 2

Figure 3. Local coordinate system for a general curved body; $\boldsymbol{e}_{\boldsymbol{z}}$ is along the tangent to the filament axis, $\boldsymbol{e}_{\boldsymbol{x}}$ is along the normal and $\boldsymbol{e}_{\boldsymbol{y}}$ is pointed along the binormal to the centreline of the slender body ($\boldsymbol{r}_{\boldsymbol{c}}$).

Figure 3

Table 1. Values of the various parameters corresponding to E. coli used in RFT calculation.

Figure 4

Figure 4. A schematic showing the inner, outer and Brinkman regions for the two-fluid model with $a \ll L_B \ll l$.

Figure 5

Figure 5. Geometry of the helix used in the numerical calculation.

Figure 6

Figure 6. Plots of normalised (a) thrust, (b) drag and (c) torque for a slender helical fibre with spheroidal cross-section translating with $U$ and rotating with $\varOmega$ in a single-fluid medium as a function of $L/R_{Helix}$ from the numerical solutions of single-fluid SBT with no-slip and slip ($\boldsymbol {f}{\cdot } \boldsymbol {e_z} = 0$). (d) Force-free swimming velocity of a rotating helix with and without slip.

Figure 7

Figure 7. A schematic of a segment of helix showing the forces acting on the segment due to rotation and translation. The thrust and torque correspond to $f_{z}$ and $f_{x}$ in the former case and drag corresponds to $f_{z}$ in the latter. Note, for simplicity, we choose $\psi = {\rm \pi} /4$ as the pitch angle of the helix, denoting the orientation of the segment with respect to the motion.

Figure 8

Figure 8. Plots of normalised (a) thrust, (b) drag and (c) torque for a slender helical fibre with spheroidal cross-section, as a function of $\lambda$, from the numerical solutions for axial translation ($U$) and rotation ($\varOmega$) of the helix in a two-fluid medium with polymer slip, where the curves correspond to different $L_B/L$. Mixture ($L_B/L \rightarrow 0$) corresponds to a single-fluid medium of viscosity $\mu _s ( 1 + \lambda )$, with the no-slip condition for both fluids. Here, $R_{Helix}/L \approx 0.052$ and $a/L \approx 0.0043$.

Figure 9

Figure 9. Plot of the ratio of thrust and drag on a helix rotating and translating in a two-fluid medium with polymer slip ($\lambda = 1$). Magenta dashed line is the ratio for the same helix rotating in a mixture ($L_B/L \rightarrow 0$) and violet dashed line is the ratio in two decoupled fluids ($L_B/L \rightarrow \infty$). Here, as in figure 8, $R_{Helix}/L \approx 0.052$ and $a/L \approx 0.0043$.

Figure 10

Figure 10. Plots of normalised (a) thrust, (b) drag and (c) torque for a slender helical fibre with spheroidal cross-section, as a function of $\lambda$, from the numerical solutions for axial translation ($U$) and rotation ($\varOmega$) of the helix in a two-fluid medium with no polymer–fibre interaction, where the curves correspond to different $L_B/L$. Here, $R_{Helix}/L \approx 0.052$ and $a/L \approx 0.0043$.

Figure 11

Figure 11. Plots of the normalised (a) thrust and (b) drag on a helix rotating and translating in a two-fluid medium with no polymer–fibre interaction ($\lambda = 1$). The magenta dashed line indicates the thrust for the same helix rotating in a mixture ($L_B/L \rightarrow 0$) and the violet dashed line indicates the thrust in the solvent ($L_B/L \rightarrow \infty$). Here, $R_{Helix}/L \approx 0.052$ and $a/L \approx 0.0043$.

Figure 12

Figure 12. (a) Plots of the ratio of the thrust and drag on a helix rotating and translating in a two-fluid medium with no polymer–fibre interaction. Single-fluid (green, dashed line) indicates the thrust to drag ratio for the same helix in both mixture ($L_B/L \rightarrow 0$) and solvent ($L_B/L \rightarrow \infty$). Note that this ratio is the same for mixture and solvent, as the ratio is independent of viscosity. (b) Plots of the ratio for small $L_B/L$. Here, $R_{Helix}/L \approx 0.052$ and $a/L \approx 0.0043$.

Figure 13

Figure 13. A swimming bacterium in a two-fluid medium with a slipping polymer at the flagellar bundle ($L_B/a_0 \gg 1$, red; mixture ($L_B/a_0 \sim 1$), magenta) and with no polymer–bundle interaction (blue) (the cell sees the mixture): (a) swimming velocity; (b) angular velocity of the cell and (c) flagellar bundle. Polymer slip ($L_B/a_0 \gg 1$) leads to a two-fold increase in swimming velocity for $\lambda > 1$.

Figure 14

Figure 14. Case of a bacterium in a two-fluid medium with a slipping polymer, where the cell sees the mixture: (a) swimming velocity; and the angular velocities of (b) bacterial head and (c) flagellar bundle for different values of $L_B/a_0$. Here, $L_B/a_0 \gg 1$, screening length being in the outer region, and mixture ($L_B/a_0 \sim 1$), screening length in the inner region, while the other curves correspond to screening length in the matching region.

Figure 15

Figure 15. Case of a bacterium in a two-fluid medium with slipping polymer ($L_B/a_0 \sim 1$): (a) swimming velocity, and the angular velocities of (b) bacterial head and (c) flagellar bundle for different values of $L_B/R_{Cell}$. Results are compared with the case where the bacterium swims in the mixture (magenta dashed curve).

Figure 16

Figure 16. Case of a bacterium in a two-fluid medium with slipping polymer ($L_B/a \gg 1$): (a) swimming velocity, and the angular velocities of (b) bacterial head and (c) flagellar bundle for different values of $L_B/R_{Cell}$. Results are compared with the case where the bacterium swims in the mixture (magenta dashed curve).

Figure 17

Figure 17. Case of a bacterium in a two-fluid medium, with non-interacting polymer (on bundle) and slipping polymer on head: (a) swimming velocity, and the angular velocities of (b) bacterial head and (c) flagellar bundle for different values of $L_B/R_{Cell}$. Results are compared with the case where the bacterium swims in the mixture (magenta dashed curve).

Figure 18

Table 2. Table summarising the key result of RFT calculations for a bacterium in a two-fluid medium, corresponding to different cases. Note that the head always moves in the mixture in the cases listed above.

Figure 19

Figure 18. Plots of the (a) angular velocity of the flagellar bundle and (b) velocity of the bacterium, normalised by the respective values in a solvent of viscosity $\mu _s$, as a function of normalised viscosity $\mu /\mu _s$, calculated using the three versions of RFT. Our two-fluid RFT (labelled TF) pertains to the case with polymer slip at the bundle ($L_B/a \gg 1$), while the head ‘sees’ a mixture. The velocity is compared with experimental measurements of Martinez et al. (2014).

Figure 20

Table 3. Values of the parameters used by Martinez et al. (2014) in RFT calculations.

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