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Stress relaxation in a dilute bacterial suspension: the active–passive transition

Published online by Cambridge University Press:  15 May 2019

Sankalp Nambiar
Affiliation:
Engineering Mechanics Unit, JNCASR, Jakkur, Bangalore 560064, India
Phanikanth S.
Affiliation:
Department of Chemical Engineering, IISc, Bangalore 560012, India
P. R. Nott
Affiliation:
Department of Chemical Engineering, IISc, Bangalore 560012, India
Ganesh Subramanian*
Affiliation:
Engineering Mechanics Unit, JNCASR, Jakkur, Bangalore 560064, India
*
†Email address for correspondence: sganesh@jncasr.ac.in

Abstract

This paper follows a recent article of Nambiar et al. (J. Fluid Mech., vol. 812, 2017, pp. 41–64) on the linear rheological response of a dilute bacterial suspension (e.g. E. coli) to impulsive starting and stopping of simple shear flow. Here, we analyse the time dependent nonlinear rheology for a pair of linear flows – simple shear (a canonical weak flow) and uniaxial extension (a canonical strong flow), again in response to impulsive initiation and cessation. The rheology is governed by the bacterium orientation distribution which satisfies a kinetic equation that includes rotation by the imposed flow, and relaxation to isotropy via rotary diffusion and tumbling. The relevant dimensionless parameters are the Péclet number $Pe\equiv \dot{\unicode[STIX]{x1D6FE}}\unicode[STIX]{x1D70F}$, which dictates the importance of flow-induced orientation anisotropy, and $\unicode[STIX]{x1D70F}D_{r}$, which quantifies the relative importance of the two intrinsic orientation decorrelation mechanisms (tumbling and rotary diffusion). Here, $\unicode[STIX]{x1D70F}$ is the mean run duration of a bacterium that exhibits a run-and-tumble dynamics, $D_{r}$ is the intrinsic rotary diffusivity of the bacterium and $\dot{\unicode[STIX]{x1D6FE}}$ is the characteristic magnitude of the imposed velocity gradient. The solution of the kinetic equation is obtained numerically using a spectral Galerkin method, that yields the rheological properties (the shear viscosity, the first and second normal stress differences for simple shear, and the extensional viscosity for uniaxial extension) over the entire range of $Pe$. For simple shear, we find that the stress relaxation predicted by our analysis at small $Pe$ is in good agreement with the experimental observations of Lopez et al. (Phys. Rev. Lett., vol. 115, 2015, 028301). However, the analysis at large $Pe$ yields relaxations that are qualitatively different. Upon step initiation of shear, the rheological response in the experiments corresponds to a transition from a nearly isotropic suspension of active swimmers at small $Pe$, to an apparently (nearly) isotropic suspension of passive rods at large $Pe$. In contrast, the computations yield the expected transition to a nearly flow-aligned suspension of passive rigid rods at high $Pe$. We probe this active–passive transition systematically, complementing the numerical solution with analytical solutions obtained from perturbation expansions about appropriate base states. Our study suggests courses for future experimental and analytical studies that will help understand relaxation phenomena in active suspensions.

JFM classification

Information

Type
JFM Papers
Copyright
© 2019 Cambridge University Press 
Figure 0

Figure 1.The iso-contours of the orientation probability density, at three different time instants, plotted for axisymmetric extension at (a) $Pe=0.1$ and (b) at $Pe=10$.

Figure 1

Figure 2.The time dependent bacterial extensional viscosity ($\langle \unicode[STIX]{x1D70E}_{E}^{b}\rangle _{33}$), given by (2.18) for step initiation of the flow at $t=0$, plotted for $Pe\sim 0.01{-}50$. The inset consists of the extensional viscosity for $Pe=10$ and $Pe=50$, with the time being scaled by the inverse rate of extension.

Figure 2

Figure 3.The iso-contours of the orientation probability density, at three different time instants, plotted for simple shear at (a) $Pe=0.1$ and (b) at $Pe=63.9$.

Figure 3

Figure 4.The time dependent shear viscosity ($\langle \unicode[STIX]{x1D70E}_{S}\rangle _{12}$), given by (2.19) for step initiation of the flow at $t=0$, plotted for $Pe\sim 0.022{-}100$. The inset consists of the shear viscosity, for $Pe=34.6$, $Pe=63.9$ and $Pe=100$, plotted as a function of the time scaled by the inverse shear rate.

Figure 4

Figure 5.The time dependent shear viscosity ($\langle \unicode[STIX]{x1D70E}_{S}\rangle _{12}$), given by (2.19) for step-down from steady flow to rest is plotted (a) for a range of $Pe\sim 0.022{-}100$. The inset has $\langle \unicode[STIX]{x1D70E}_{S}\rangle _{12}$ scaled by $Pe^{4/3}$ for $Pe=0.138,63.9,100$. The collapse of the largest two $Pe$ is consistent with the scaling of the active stress in the $Pe\gg 1$ regime. (b) Comparison of the $\unicode[STIX]{x1D6E4}$ distributed relaxation with a shape parameter $k=0.64$ that best fits an exponential relaxation of $2\unicode[STIX]{x1D70F}$ (Nambiar et al.2017) for $Pe=0.022$ and 5.

Figure 5

Figure 6.The time dependent shear viscosity (a) reported in Lopez et al. (2015) (reproduced with permission – © 2015 American Physical Society), and (b) given by (2.19), with the solvent contribution included, plotted for $Pe\sim 0.022{-}63.9$. For E. coli, the run duration is $\unicode[STIX]{x1D70F}\approx 1~\text{s}$, and hence $Pe$ equals the shear rate $\dot{\unicode[STIX]{x1D6FE}}$ (in $\text{s}^{-1}$).

Figure 6

Figure 7.The time dependent bacterial normal stress differences determined numerically in (2.20a) and (2.20b) are plotted for $Pe\sim [0.01,100]$. (a) $N_{1}$ (b) $N_{2}$.

Figure 7

Figure 8.Comparison of the numerical and analytical high-$Pe$ relaxations. (a) Axisymmetric extensional flow, (b) simple shear flow at $Pe_{r}=100$, 200 and 300. The solid line represent the analytically obtained stress, equations (4.6) and (4.10), respectively, while the dashed, dotted and dash-dotted lines represent the numerical solutions.

Figure 8

Figure 9.Comparison of steady-state extensional viscosity with Saintillan (2010b) for two values of dipole strength that were chosen by them. The dashed lines represent data obtained from Saintillan (2010b), while the markers (♢ and ○) represent our data.

Figure 9

Figure 10.Comparison of steady-state rheological properties with Saintillan (2010a): (a) shear viscosity, (b) first normal stress difference ($N_{1}$) and (c) second normal stress difference ($N_{2}$). The dashed lines represent data obtained from Saintillan (2010a), while the markers (♢, ○) represent our data.

Figure 10

Figure 11.Comparison of absolute values of the steady-state (a) first normal stress difference ($N_{1}$) and (b) second normal stress difference ($N_{2}$), for varying dipole strength. The ♢ marker correspond to the dipole strength we consider in the main text (and this is representative of the experiments of Lopez et al. (2015)) and the ○ marker corresponds to $1/10$th of the same. The discontinuity in (a) for the reduced dipole strength (○ marker) is indicative of a sign change in $N_{1}$.

Figure 11

Figure 12.Comparison of the absolute values of the active and passive contributions at steady-state (a) first normal stress difference ($N_{1}$) and (b) second normal stress difference ($N_{2}$), for varying dipole strength. The ♢ and ○ markers correspond to the contribution from the active component for the respective dipole strengths of 1.8 and 0.18. The ▫ markers corresponds to the contribution from the passive component.

Figure 12

Figure 13.Comparison of time dependent responses of bacterial viscosities at low $Pe$ obtained numerically with the (a) Extensional viscosity, with the analytical results evaluated to $O(Pe^{2})$, and (b) shear viscosity, with the analytical results evaluated to $O(Pe^{3})$. The dashed lines represent the analytically obtained viscosities, while the solid lines correspond to the viscosities obtained numerically.

Figure 13

Figure 14.Comparison of time dependent responses of bacterial normal stress differences at low $Pe$ obtained from the $O(Pe^{2})$ analysis with numerics for the case of simple shear flow. (a) $N_{1}$ (b) $N_{2}$. The dashed lines represent the analytically obtained viscosities, while the solid lines correspond to the viscosities obtained numerically.

Figure 14

Figure 15.Comparison of steady-state orientation probability density function ($\lim _{t\rightarrow \infty }\unicode[STIX]{x1D6FA}(\boldsymbol{p},t)$) along the flow-gradient plane for simple shear flow (a) $Pe=0.1$, (b) $Pe=100$.

Figure 15

Figure 16.Comparison of time dependent viscosity for different set of terms in the series expansion. The dotted curve (brown), the straight curve (light maroon) and the dashed curve (blue), correspond to considering (a) 20, 48 and 58 terms for axisymmetric extensional flow and (b) 121, 441 and 961 terms for simple shear flow, respectively in the series expansion of $\unicode[STIX]{x1D6FA}(\boldsymbol{p},t)$.