Hostname: page-component-74656f9884-8r8nv Total loading time: 0 Render date: 2026-11-10T05:03:06.244Z Has data issue: false hasContentIssue false

Viscous propulsion in active transversely isotropic media

Published online by Cambridge University Press:  05 January 2017

G. Cupples
Affiliation:
School of Mathematics, University of Birmingham, Birmingham B15 2TT, UK
R. J. Dyson
Affiliation:
School of Mathematics, University of Birmingham, Birmingham B15 2TT, UK
D. J. Smith*
Affiliation:
School of Mathematics, University of Birmingham, Birmingham B15 2TT, UK Institute for Metabolism and Systems Research, University of Birmingham, Birmingham B15 2TT, UK Centre for Human Reproductive Science, Birmingham Women’s NHS Foundation Trust, Birmingham B15 2TG, UK
*
†Email address for correspondence: D.J.Smith.2@bham.ac.uk

Abstract

Taylor’s swimming sheet is a classical model of microscale propulsion and pumping. Many biological fluids and substances are fibrous, having a preferred direction in their microstructure; for example, cervical mucus is formed of polymer molecules which create an oriented fibrous network. Moreover, suspensions of elongated motile cells produce a form of active oriented matter. To understand how these effects modify viscous propulsion, we extend Taylor’s classical model of small-amplitude zero-Reynolds-number propulsion of a ‘swimming sheet’ via the transversely isotropic fluid model of Ericksen, which is linear in strain rate and possesses a distinguished direction. The energetic costs of swimming are significantly altered by all rheological parameters and the initial fibre angle. Propulsion in a passive transversely isotropic fluid produces an enhanced mean rate of working, independent of the initial fibre orientation, with an approximately linear dependence of the energetic cost on the extensional and shear enhancements to the viscosity caused by fibres. In this regime, the mean swimming velocity is unchanged from the Newtonian case. The effect of the constant term in Ericksen’s model for the stress, which can be identified as a fibre tension or alternatively a stresslet characterising an active fluid, is also considered. This stress introduces an angular dependence and dramatically changes the streamlines and flow field; fibres aligned with the swimming direction increase the energetic demands of the sheet. The constant fibre stress may result in a reversal of the mean swimming velocity and a negative mean rate of working if it is sufficiently large relative to the other rheological parameters.

Information

Type
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 in any medium, provided the original work is properly cited.
Copyright
© 2017 Cambridge University Press
Figure 0

Figure 1. Parallel filament mesh in cervical mucus during the time of ovulation; $\text{bar}=10~\unicode[STIX]{x03BC}\text{m}$. Republished with permission of Oxford University Press, from ‘Ultrastructure of the Human Periovulatory Cervical Mucus’, Ceric et al. (2005); permission conveyed through Copyright Clearance Center, Inc.

Figure 1

Figure 2. A schematic of Taylor’s swimming sheet in a Newtonian fluid. Working in a frame of reference in which the sheet is stationary, $b^{\ast }$ is the amplitude, $\unicode[STIX]{x1D706}^{\ast }=2\unicode[STIX]{x03C0}/k^{\ast }$ is the wavelength and $k^{\ast }$ is the wavenumber. The flow at infinity in the $x^{\ast }$-direction is $U^{\ast }$. A travelling wave traverses the sheet with speed $c^{\ast }=\unicode[STIX]{x1D714}^{\ast }/k^{\ast }$, where $\unicode[STIX]{x1D714}^{\ast }$ is the angular velocity.

Figure 2

Figure 3. Regimes of interest in the parameter space: (i) the plane $\unicode[STIX]{x1D707}_{1}=0$ is the passive transversely isotropic regime; (ii) the dashed line represents the active-only regime where $\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=0$; (iii) the quarter cylinder is the nearly isotropic regime where neither $\unicode[STIX]{x1D707}_{2}$ nor $\unicode[STIX]{x1D707}_{3}$ is large; (iv) the remaining region is the fully active and transversely isotropic regime.

Figure 3

Figure 4. A schematic showing the initial uniform orientation angle, $\unicode[STIX]{x1D719}$, and the small perturbation away from this angle, $\unicode[STIX]{x1D703}$.

Figure 4

Figure 5. Mean rate of working by the swimmer in a transversely isotropic fluid where $\unicode[STIX]{x1D707}_{1}=0$. (a) The mean rate of working for varying $\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$. This result is identical for all initial angles $\unicode[STIX]{x1D719}$. (b) The change in the mean rate of working for increasing $\unicode[STIX]{x1D707}_{\Vert }$ for a range of $\unicode[STIX]{x1D707}_{3}$ and set $\unicode[STIX]{x1D707}_{2}$ values: $\unicode[STIX]{x1D707}_{2}=0$ (solid line), $\unicode[STIX]{x1D707}_{2}=300$ (dashed line), $\unicode[STIX]{x1D707}_{2}=600$ (dash-dotted line) and $\unicode[STIX]{x1D707}_{2}=900$ (dotted line).

Figure 5

Figure 6. Mean swimming velocity versus the initial orientation $\unicode[STIX]{x1D719}$, where $\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=0$. Panels (a) and (b) depict small positive and negative $\unicode[STIX]{x1D707}_{1}$ values: $0$ (solid line), $\pm 0.1$ (dashed line), $\pm 1$ (dash-dotted line) and $\pm 5$ (dotted line). Panels (c) and (d) depict larger $\unicode[STIX]{x1D707}_{1}$ values: $0$ (solid line), $\pm 300$ (dashed line), $\pm 600$ (dash-dotted line), $\pm 900$ (dotted line), where the arrows denote increasing $\unicode[STIX]{x1D707}_{1}$.

Figure 6

Figure 7. Mean rate of working versus the initial orientation $\unicode[STIX]{x1D719}$, where $\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=0$. Panels (a) and (b) depict small positive and negative $\unicode[STIX]{x1D707}_{1}$ values: $0$ (solid line), $\pm 0.1$ (dashed line), $\pm 1$ (dash-dotted line) and $\pm 5$ (dotted line). Panels (c) and (d) depict larger $\unicode[STIX]{x1D707}_{1}$ values: $0$ (solid line), $\pm 300$ (dashed line), $\pm 600$ (dash-dotted line), $\pm 900$ (dotted line), where the arrows denote increasing $\unicode[STIX]{x1D707}_{1}$.

Figure 7

Figure 8. Mean swimming velocity versus $\unicode[STIX]{x1D719}$, where the parameters $\unicode[STIX]{x1D707}_{1},\,\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$ take values of up to $5$. Panels (a) and (b) show $\unicode[STIX]{x1D707}_{1}=\pm 0.1$, (c) and (d) show $\unicode[STIX]{x1D707}_{1}=\pm 1$ and (e) and (f) show $\unicode[STIX]{x1D707}_{1}=\pm 5$. Each line depicts a different $\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$ combination: $\unicode[STIX]{x1D707}_{2}=0,\,\unicode[STIX]{x1D707}_{3}=0$ (solid line), $\unicode[STIX]{x1D707}_{2}=1,\,\unicode[STIX]{x1D707}_{3}=0$ (dashed line), $\unicode[STIX]{x1D707}_{2}=0,\,\unicode[STIX]{x1D707}_{3}=1$ (dash-dotted line) and $\unicode[STIX]{x1D707}_{2}=1,\,\unicode[STIX]{x1D707}_{3}=1$ (dotted line).

Figure 8

Figure 9. Mean rate of working versus $\unicode[STIX]{x1D719}$, where the parameters $\unicode[STIX]{x1D707}_{1},\,\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$ take values of up to $5$. Panels (a) and (b) show $\unicode[STIX]{x1D707}_{1}=\pm 0.1$, (c) and (d) show $\unicode[STIX]{x1D707}_{1}=\pm 1$ and (e) and (f) show $\unicode[STIX]{x1D707}_{1}=\pm 5$. Each line depicts a different $\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$ combination: $\unicode[STIX]{x1D707}_{2}=0,\,\unicode[STIX]{x1D707}_{3}=0$ (solid line), $\unicode[STIX]{x1D707}_{2}=1,\,\unicode[STIX]{x1D707}_{3}=0$ (dashed line), $\unicode[STIX]{x1D707}_{2}=0,\,\unicode[STIX]{x1D707}_{3}=1$ (dash-dotted line) and $\unicode[STIX]{x1D707}_{2}=1,\,\unicode[STIX]{x1D707}_{3}=1$ (dotted line).

Figure 9

Figure 10. Mean swimming velocity versus $\unicode[STIX]{x1D719}$ when at least one of $\unicode[STIX]{x1D707}_{1},\,\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$ is much larger than 1. Panels (a) and (c) depict positive $\unicode[STIX]{x1D707}_{1}$ values and (b) and (d) depict negative $\unicode[STIX]{x1D707}_{1}$ values. The values $\unicode[STIX]{x1D707}_{1}$ takes are $0$ (solid line), $\pm 300$ (dashed line), $\pm 600$ (dash-dotted line) and $\pm 900$ (dotted line). In (a) and (b), $\unicode[STIX]{x1D707}_{2}=900,\,\unicode[STIX]{x1D707}_{3}=0$, and in (c) and (d), $\unicode[STIX]{x1D707}_{2}=0,\,\unicode[STIX]{x1D707}_{3}=900$, where the arrows denote increasing $\unicode[STIX]{x1D707}_{1}$.

Figure 10

Figure 11. Mean rate of working versus $\unicode[STIX]{x1D719}$ when at least one of $\unicode[STIX]{x1D707}_{1},\unicode[STIX]{x1D707}_{2}$ and $\unicode[STIX]{x1D707}_{3}$ is much larger than 1. Panels (a) and (c) depict positive $\unicode[STIX]{x1D707}_{1}$ values and (b) and (d) depict negative $\unicode[STIX]{x1D707}_{1}$ values. The values $\unicode[STIX]{x1D707}_{1}$ takes are $0$ (solid line), $\pm 300$ (dashed line), $\pm 600$ (dash-dotted line) and $\pm 900$ (dotted line). In (a) and (b), $\unicode[STIX]{x1D707}_{2}=900,\unicode[STIX]{x1D707}_{3}=0$, and in (c) and (d), $\unicode[STIX]{x1D707}_{2}=0,\unicode[STIX]{x1D707}_{3}=900$, where the arrows denote increasing $\unicode[STIX]{x1D707}_{1}$.

Figure 11

Figure 12. Fibre angle, $\unicode[STIX]{x1D719}+\unicode[STIX]{x1D700}\unicode[STIX]{x1D703}_{0}$, in passive and active regimes: (a) the passive regime ($\unicode[STIX]{x1D707}_{1}=0,\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=5$) and (b) the active-only regime ($\unicode[STIX]{x1D707}_{1}=5,\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=0$). In each panel, $t=0$, $\unicode[STIX]{x1D700}=0.2$ and the initial orientation angle is $\unicode[STIX]{x1D719}=0$. See movie 1.

Figure 12

Figure 13. Velocity field in passive and active regimes: (a) passive regime ($\unicode[STIX]{x1D707}_{1}=0,\,\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=5$) and (b) active-only regime ($\unicode[STIX]{x1D707}_{1}=5,\,\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=0$). In each panel, $t=0$ and the initial orientation angle is $\unicode[STIX]{x1D719}=0$. See movie 2.

Figure 13

Figure 14. Instantaneous streamlines in the four regimes considered: (a) passive regime ($\unicode[STIX]{x1D707}_{1}=0,\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=5$), (b) active-only regime ($\unicode[STIX]{x1D707}_{1}=5,\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=0$), (c) nearly isotropic regime ($\unicode[STIX]{x1D707}_{1}=\unicode[STIX]{x1D707}_{3}=1,\unicode[STIX]{x1D707}_{2}=0$) and (d) regime where at least one of the parameters is much larger than 1 ($\unicode[STIX]{x1D707}_{1}=\unicode[STIX]{x1D707}_{2}=\unicode[STIX]{x1D707}_{3}=900$). In each panel, $t=0$ and the initial orientation angle is $\unicode[STIX]{x1D719}=0$. See movie 3.

Cupples et al. supplementary movie

Movie visualisation of figure 12, depicting time-evolution of fibre angle above the swimming sheet. (a) passive regime ('_1=0, '_2='_3=5), (b) active-only regime ('_1=5, '_2='_3=0), (c) nearly-isotropic regime ('_1='_3=1, '_2=0) and (d) regime where at least one of the parameters is much larger than one ('_1='_2='_3=900). In each graph t=0, e=0.2 and the initial orientation angle is f=0.

Download Cupples et al. supplementary movie(Video)
Video 1.7 MB

Cupples et al. supplementary movie

Movie visualisation of figures 13, depicting time-evolution of velocity field above the swimming sheet. (a) passive regime ('_1=0, '_2='_3=5), (b) active-only regime ('_1=5, '_2='_3=0), (c) nearly-isotropic regime ('_1='_3=1, '_2=0) and (d) regime where at least one of the parameters is much larger than one ('_1='_2='_3=900). In each graph t=0, e=0.2 and the initial orientation angle is f=0.

Download Cupples et al. supplementary movie(Video)
Video 8.4 MB

Cupples et al. supplementary movie

Movie visualisation of figures 14, depicting time-evolution of streamlines above the swimming sheet. (a) passive regime ('_1=0, '_2='_3=5), (b) active-only regime ('_1=5, '_2='_3=0), (c) nearly-isotropic regime ('_1='_3=1, '_2=0) and (d) regime where at least one of the parameters is much larger than one ('_1='_2='_3=900). In each graph t=0, e=0.2 and the initial orientation angle is f=0.

Download Cupples et al. supplementary movie(Video)
Video 8 MB