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Intermittent swimmers optimize energy expenditure with flick-to-flick motor control

Published online by Cambridge University Press:  10 March 2025

Yi Zhu
Affiliation:
Key Laboratory of Coastal Environment and Resources of Zhejiang Province, School of Engineering, Westlake University, Hangzhou, Zhejiang 310030, PR China
Linlin Kang*
Affiliation:
Key Laboratory of Coastal Environment and Resources of Zhejiang Province, School of Engineering, Westlake University, Hangzhou, Zhejiang 310030, PR China
Xinyu Tong
Affiliation:
Key Laboratory of Coastal Environment and Resources of Zhejiang Province, School of Engineering, Westlake University, Hangzhou, Zhejiang 310030, PR China
Jingtao Ma
Affiliation:
Aix Marseille Univ, CNRS, Centrale Marseille, M2P2, Marseille, FranceSchool of Engineering and Information Technology, University of New South Wales, Canberra, ACT 2600, Australia
Fangbao Tian*
Affiliation:
School of Engineering and Information Technology, University of New South Wales, Canberra, ACT 2600, Australia
Dixia Fan*
Affiliation:
Key Laboratory of Coastal Environment and Resources of Zhejiang Province, School of Engineering, Westlake University, Hangzhou, Zhejiang 310030, PR China
*
Corresponding authors: Linlin Kang, kanglinlin@westlake.edu.cn; Fangbao Tian, f.tian@adfa.edu.au; Dixia Fan, fandixia@westlake.edu.cn
Corresponding authors: Linlin Kang, kanglinlin@westlake.edu.cn; Fangbao Tian, f.tian@adfa.edu.au; Dixia Fan, fandixia@westlake.edu.cn
Corresponding authors: Linlin Kang, kanglinlin@westlake.edu.cn; Fangbao Tian, f.tian@adfa.edu.au; Dixia Fan, fandixia@westlake.edu.cn

Abstract

Intermittent swimming behaviour is commonly observed in larval zebrafish, often attributed to energy-saving mechanisms. In this study, we utilize a hybrid approach combining deep reinforcement learning and the immersed boundary–lattice Boltzmann method to train a larval zebrafish-like swimmer to reach a target with minimized energy expenditure. We find that when the tail-beat period is fixed, continuous swimming emerges as the optimal strategy. However, when the tail-beat period is allowed to vary, intermittent swimming proves superior in energy performance, achieved through reductions in tail-beat amplitude and frequency. Our detailed analysis reveals that intermittent swimmers employ rapid backward tail flicks to attain high speeds, coupled with slower forward tail flicks and coasting phases to conserve energy. Furthermore, we derive scaling laws governing the swimming performance of trained fish. These results offer valuable insights into the intermittent swimming patterns of fish, with implications for understanding bio-inspired locomotion and informing the design of energy-efficient aquatic systems.

Information

Type
JFM Papers
Copyright
© The Author(s), 2025. Published by Cambridge University Press
Figure 0

Figure 1.A schematic of the fish model: (a) ventral view of a larval zebrafish (Yao et al.2018); (b) constructed shape of the two-dimensional model; (c) illustration of the motion of the swimmer. Here, $L$ is the length of the fish, $l$ is the arc length along the mid-line of the body, $(x,y)$ represents the global coordinate system, while $(x_l, y_l)$ signifies the local coordinate system, whose origin is positioned at the head of the fish, $d$ is the half-thickness of the body, $r_{tip}$ is the distance between the head of the fish and the target, $\theta _{tip}$ is the orientation angle of the swimmer with respected to the target, ${\boldsymbol{r}}_c$ and $\theta$ are the position vector and the body rotation angle around the mass centre, respectively, ${\boldsymbol{F}}_t$ and $M_z$ are the total hydrodynamic force on the fish body and the total hydrodynamic torque on the centre of mass, respectively, and $h_l$ and $\theta _l$ are the lateral displacement and deflection angle of the mid-line with respect to the $x_l$-axis, respectively.

Figure 1

Figure 2.Baseline grids used in the simulation: (a) full and (b) partial computational domains.

Figure 2

Figure 3.The reinforcement learning framework in fish swimming towards a target. The agent is the DRQN; the environment is the fluid–structure interaction simulation using the IB–LBM; the action is the adjustment of swimming motion; the reward is to reduce the energy expenditure; states are distance and orientation angle to the target, and action.

Figure 3

Figure 4.The changing history of $COT$ with respective to the training episode for (a) a swimmer with FTP swimming, and (b) a swimmer with VTP swimming. The black dashed line in each plot is obtained through mean filtering of the data from the closest 100 episodes.

Figure 4

Figure 5.Trailing-edge trajectories of swimmers with FTP and VTP, and their positions and gestures at several instants with three initial settings: (a) $r_{tip,\textrm {0}}=8L$ and $\theta _{tip, \textrm {0}}=10^\circ$, (b) $r_{tip,\textrm {0}}=8L$ and $\theta _{tip, \textrm {0}}=90^\circ$, (c) $r_{tip,\textrm {0}}=3L$ and $\theta _{tip, \textrm {0}}=170^\circ$.

Figure 5

Figure 6.The temporal evolution of (a) lateral amplitude of the tail tip $y_{l,tail}$, (b) orientation angle of the head $\theta _{tip}$, (c) swimming speed $u_r$, and (d) input work coefficient $C_P$, for the fish swimming system with $r_{tip,\textrm {0}}=8L$ and $\theta _{tip,\textrm {0}}=90^\circ$. A zoomed-in view of the area located between the green vertical lines in (a) will be presented in figure 9(a). Here, $T$ is the fixed characteristic time ensuring $Re=L^2/T\nu =1000$.

Figure 6

Figure 7.Parameter space for (a–d) VTP intermittent swimming and (e–h) FTP continuous swimming. Contour plot of time-averaged (a,e) swimming speed ($\bar {U}$), (b,f) peak-to-peak amplitude of tail tip ($\bar {A}_{tail}$), (c,g) swimming period ($\bar {T}_a$) and (d,h) cost of transport ($COT$) against initial distance ($r_{tip,\textrm {0}}$) and initial oriented angle ($\theta _{tip,\textrm {0}}$), where the unit of the vertical axis is degrees.

Figure 7

Figure 8.Comparison of swimming performance of VTP intermittent swimming and FTP continuous swimming. (a) Relative deviation in cost of transport ($\triangle COT^*$) versus relative deviation in time-averaged swimming speed ($\triangle \bar {U}^*$). (b) Relative deviation in time-averaged peak-to-peak tail tip amplitude ($\triangle \bar {A}_{tail}^*$) versus relative deviation in swimming period ($\triangle \bar {T}_a^*$).

Figure 8

Figure 9.The temporal evolution of (a) lateral position of the tail tip $y_{l,tail}$, (b) swimming speed $u_r$, (c) force coefficient in the $x_l$-direction $C_F$, and (d) input power coefficient $C_P$, for the fish swimming system with $r_{tip,\textrm {0}}=8L$ and $\theta _{tip,\textrm {0}}=90^\circ$ in a short period (the time period is identified by the two green dashed lines in figure 6a). The FTP continuous swimming and VTP intermittent swimming are marked by blue dashed lines and red solid lines, respectively. In (a), black circles and green squares denote special moments from figures 10 and 11, respectively.

Figure 9

Figure 10.The instantaneous vorticity contour, and distribution of key physical variables on the body surface: $1$ velocity ${\boldsymbol{u}}_B$ (the red arrow represents the velocity vector); $2$ hydrodynamic force ${\boldsymbol{F}}_f$ (the blue arrow represents the hydrodynamic force distribution on the body); $3$ input power $P_B$, for the FTP continuous swimmer at (a) $t=25.6$, (b) $t=25.8$, (c) $t=26.0$ and (d) $t=26.3$. The lateral amplitudes of the tail tip for these special moments are marked by black circles in figure 9(a).

Figure 10

Figure 11.The instantaneous vorticity contour, and distribution of key physical variables on the body surface: $1$ velocity ${\boldsymbol{u}}_B$ (the red arrow represents the velocity vector); $2$ hydrodynamic force ${\boldsymbol{F}}_f$ (the blue arrow represents the hydrodynamic force distribution on the body); $3$ input power $P_B$, for the VTP swimmer at (a) $t=25.7$, (b) $t=26.4$, (c) $t=26.5$, (d) $t=26.6$, (e) $t=26.8$ and (f) $t=27.1$. The lateral amplitudes of the tail tip for these special moments are marked by green squares in figure 9(a).

Figure 11

Figure 12.Schematic diagram for (a) force decomposition and (b) thrust and drag contribution from pressure force and viscous force. Here, $\boldsymbol {n}$ and $\boldsymbol {\tau}$ represent the local normal and tangential vectors, respectively, ${\boldsymbol{F}}_{f,i}$ signifies the hydrodynamic force, and${\boldsymbol{F}}_i^n=(F_{x,i}^n, F_{y,i}^n)$ and ${\boldsymbol{F}}_i^{\tau }=(F_{x,i}^\tau , F_{y,i}^\tau )$ represent pressure force and viscous force acting on the $i$th surface element of the fish body. (c,d) The $x$-component of the hydrodynamic force $F_x={\boldsymbol{F}}_f\boldsymbol{\cdot} {\boldsymbol {e}}_x$, thrust $F_T$, and drag $F_D$ as functions of time for (c) FTP swimming and (d) VTP swimming. Here, ${\boldsymbol {e}}_x$ denotes the unit vector aligned with the $x$-axis.

Figure 12

Figure 13.Scaling swimming performance for swimmers with FTP (black circles) and VTP (red squares). The cruising Reynolds number $Re_u$ is a function of $Sw^*=Sw/\sqrt {(\beta -1)DC^2+DC}$, and $\beta =2.5$.

Figure 13

Figure 14.Scaling swimming performance for swimmers with FTP (black circles) and VTP (red squares). (a) The thrust $\bar {F}_T$ as a function of the modified swimming number $Sw'=Sw/DC^{0.5}$. (b) The drag $\bar {F}_D$ as a function of the modified Reynolds number $Re'_{\!\!u}=Re_u/[(\beta -1)DC+1]^{2/3}$. Here, $\beta =2.5$.

Figure 14

Figure 15.Scaling swimming performance for swimmers with FTP (black circles) and VTP (red squares). Cost of transport $COT$ is a function of $Sw^{\circ }=Sw*[(\beta -1)/DC+1/DC^2]^{2/5}$, and $\beta =2.5$.

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