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Morphology and energetics of the wake behind a continuously swimming crucian carp at different flow velocities


Abstract

Understanding the hydrodynamic principles underlying fish swimming efficiency is critical for optimizing fishway design and improving habitat restoration. Wake morphology and energetics of continuously swimming crucian carp were examined across flow velocities in a Brett-type swimming tunnel and analyzed using the velocity gradient method. High-speed video captured swimming behavior and two-dimensional particle image velocimetry were used to visualize the flow field generated by the fish. The critical swimming speed (Ucrit) of crucian carp was 0.85 m/s and wake vortices were observed across flow velocity gradients ranging from 0.15 to 1.2 m/s. The wake beat frequency and vorticity of the wake increasing linearly with flow velocity. Hydrodynamic efficiency of the wake vortex energy ranged from 62 to 84%. Significant changes in tailbeat amplitude, head beat amplitude, and stride length were observed near (70–88%) Ucrit (0.60–0.75 m/s), when the dimensionless Strouhal number (St) indicates efficient swimming. Thus, we conclude that 70%-88% of Ucrit is the threshold range for efficient swimming and this provides practical guidance for setting flow velocities in fishways and habitat restoration projects that will enhance fish survival.

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Data availability

The datasets used and/or analyzed in the current study are available from the corresponding author on reasonable request.

References

  1. Lauder, G. V. & Drucker, E. G. Forces, fishes, and fluids: Hydrodynamic mechanisms of aquatic locomotion. News Physiol. Sci. 17, 235–240 (2002).

    Google Scholar 

  2. Liao, J. C. A review of fish swimming mechanics and behaviour in altered flows. Philos. Trans. Roy. Soc. B-Biol. Sci. 362(1487), 1973–1993 (2007).

    Google Scholar 

  3. Krueger, Y., Hanke, W., Miersch, L. & Dehnhardt, G. Detection and direction discrimination of single vortex rings by harbour seals (Phoca vitulina). J. Exp. Biol. 221(8), jeb170753 (2018).

    Google Scholar 

  4. Mueller, U. K., van den Heuvel, B. L. E., Stamhuis, E. & Videler, J. J. Fish foot prints: Morphology and energetics of the wake behind a continuously swimming mullet (Chelon labrosus risso). J. Exp. Biol. 200(22), 2893–2906 (1997).

    Google Scholar 

  5. Muller, U. K., Smit, J., Stamhuis, E. J. & Videler, J. J. How the body contributes to the wake in undulatory fish swimming: Flow fields of a swimming eel (Anguilla anguilla). J. Exp. Biol. 204(16), 2751–2762 (2001).

    Google Scholar 

  6. Link, O. et al. The fish Strouhal number as a criterion for hydraulic fishway design. Ecol. Eng. 103, 118–126 (2017).

    Google Scholar 

  7. Willert, C. E. & Gharib, M. Digital particle image velocimetry. Exp. Fluids 10(4), 181–193 (1991).

    Google Scholar 

  8. Anderson, E. J., Mcgillis, W. R. & Grosenbaugh, M. A. The boundary layer of swimming fish. J. Exp. Biol. 204(Pt 1), 81–102 (2001).

    Google Scholar 

  9. Bartol, I. K: Hydrodynamic stability of swimming in ostraciid fishes: Role of the carapace in the smooth trunkfish Lactophrys triqueter (Teleostei: Ostraciidae). J. Exp. Biol. 206(4), 725–744 (2003).

    Google Scholar 

  10. Plaut, I. Critical swimming speed: its ecological relevance. Comp. Biochem. Physiol. A: Mol. Integr. Physiol. 131(1), 41–50 (2001).

    Google Scholar 

  11. Cano-Barbacil, C. et al. Key factors explaining critical swimming speed in freshwater fish: A review and statistical analysis for Iberian species. Sci. Rep. 10(1), 18947 (2020).

    Google Scholar 

  12. Tu, Z. et al. Aerobic swimming performance of juvenile Schizothorax chongi (Pisces, Cyprinidae) in the Yalong River, southwestern China. Hydrobiologia 675(1), 119–127 (2011).

    Google Scholar 

  13. Cai, L. et al. Swimming capability and swimming behavior of juvenile Acipenser schrenckii. J. Exp. Zool. Part a-Ecol. Genet. Physiol. 319A(3), 149–155 (2013).

    Google Scholar 

  14. Ohlberger, J., Staaks, G. & Holker, F. Swimming efficiency and the influence of morphology on swimming costs in fishes. J. Comp. Physiol. B Biochem. Syst. Environ. Physiol. 176(1), 17 (2006).

    Google Scholar 

  15. Zhang, S.-l, Zhang, J.-b, Qiao, Y., Wang, L.-t & Hou, Y.-q. Experiment study on aerobic swimming performance and behavior of Schizothorax wangchiachii Fang. J. Hydroecol. 37(005), 56–62 (2016).

    Google Scholar 

  16. Romano, D., Wahi, A., Miraglia, M. & Stefanini, C. Development of a novel underactuated robotic fish with magnetic transmission system. Machines 10, 755 (2022).

    Google Scholar 

  17. Manduca, G., Padovani, L., Carosio, E., Graziani, G., Stefanini, C., Romano, D. Development of an autonomous fish-inspired robotic platform for aquaculture inspection and management. In 2023 IEEE International Workshop on Metrology for Agriculture and Forestry (MetroAgriFor) 6–8, 188–193 (2023).

  18. Manduca, G., Santaera, G., Dario, P., Stefanini, C. & Romano, D. IEEE: How to achieve maneuverability and adaptability in an underactuated robotic fish by using a bio-inspired control approach. In IEEE/RSJ International Conference on Intelligent Robots and Systems (IROS): 2023 Oct 01–05 2023, Detroit, MI, 4757–4762 (2023).

  19. Berg, S. C. V. D., Scharff, R. B. N., Rusák, Z. & Wu, J. OpenFish: Biomimetic design of a soft robotic fish for high speed locomotion. J HardwareX 12, e00320 (2022).

    Google Scholar 

  20. Lin, Y. H., Siddall, R., Schwab, F., Fukushima, T. & Jusufi, A. Modeling and control of a soft robotic fish with integrated soft sensing. Adv. Intell. Syst. 5, 2000244 (2021).

    Google Scholar 

  21. He, F. et al. A method for estimating the velocity at which anaerobic metabolism begins in swimming fish. Water 13(10), 1430 (2021).

    Google Scholar 

  22. Dicicco, R. M. et al. Retinal regeneration following OCT-guided laser injury in zebrafish. Investig. Ophthalmol. Vis. Sci. 55(10), 6281–6288 (2014).

    Google Scholar 

  23. Muller, U. K., VandenHeuvel, B. L. E., Stamhuis, E. J. & Videler, J. J. Fish foot prints: Morphology and energetics of the wake behind a continuously swimming mullet (Chelon labrosus risso). J. Exp. Biol. 200(22), 2893–2906 (1997).

    Google Scholar 

  24. Spedding, G. R., Rayner, J. M. V. & Pennycuick, C. J. Momentum and energy in the wake of a pigeon (Columba livia) in slow flight. J. Exp. Biol. 111(1), 81–102 (1984).

    Google Scholar 

  25. Sg, R. The wake of a kestrel (Falco tinnunculus) in flapping flight. J. Exp. Biol. 127, 59–78 (1987).

    Google Scholar 

  26. Vogel, S. Life in Moving Fluids (Priceton University Press, Princeton, New Jersey, 1994).

    Google Scholar 

  27. Gu, K.-H., Chang, K.-H. & Chang, T.-J. Numerical investigation on three-dimensional fishway hydrodynamics and fish passage energetics. J. Taiwan Agric. Eng. 54(3), 64–84 (2007).

    Google Scholar 

  28. Khan, L. A. A three-dimensional computational fluid dynamics (CFD) model analysis of free surface hydrodynamics and fish passage energetics in a vertical-slot fishway. N. Am. J. Fish. Manag. 26(2), 255–267 (2006).

    Google Scholar 

  29. Haefner, J. W. & Bowen, M. D. Physical-based model of fish movement in fish extraction facilities. Ecol. Model. 152(2–3), 227–245 (2002).

    Google Scholar 

  30. Rosen, M. W. Water Flow About a Swimming Fish (1959).

  31. Lighthill, M. J. Aquatic propulsion of high hydrodynamic efficiency. J Fluid Mech 44, 265–301 (1970).

    Google Scholar 

  32. Ahlborn, B., Harper, D. G., Blake, R. W., Ahlborn, D. & Cam, M. Fish without footprints. J. Theor. Biol. 148(4), 521–533 (1991).

    Google Scholar 

  33. Videler, J. J. Fish Swimming (Springer, Netherlands, 1993).

    Google Scholar 

  34. Fish, F. E. & Lauder, G. V. Passive and active flow control by swimming fishes and mammals. Annu. Rev. Fluid Mech. 38, 193–224 (2006).

    Google Scholar 

  35. Kai, Z. Study on propulsion and energy absorption mechanism of typical underwater bionic foils. PhD. Harbin Institute of Technology (2017).

  36. Zhu, Y.-P., Cao, Z.-D. & Fu, S.-J. Aerobic and anaerobic metabolism in response to different swimming speed of juvenile darkbarbel catfish (pelteobagrus vachelli richardson). Acta Hydrobiol. Sin. 34(5), 905–912 (2010).

    Google Scholar 

  37. Burgetz, I. J., Rojas-Vargas, A., Hinch, S. G. & Randall, D. J. Initial recruitment of anaerobic metabolism during sub-maximal swimming in rainbow trout (Oncorhynchus mykiss). J. Exp. Biol. 201(19), 2711–2721 (1998).

    Google Scholar 

  38. Wardle, C. S., Videler, J. J. & Altringham, J. D. Tuning in to fish swimming waves: body form, swimming mode and muscle function. J. Exp. Biol. 198(Pt 8), 1629–1636 (1995).

    Google Scholar 

  39. Brett, J. R. The respiratory metabolism and swimming performance of young sockeye salmon. J. Fish. Res. Board Canada 21, 1183 (1964).

    Google Scholar 

  40. Lee, C. G., Farrell, A. P., Lotto, A., Hinch, S. G. & Healey, M. C. Excess post-exercise oxygen consumption in adult sockeye (Oncorhynchus nerka) and coho (O-kisutch) salmon following critical speed swimming. J. Exp. Biol. 206(18), 3253–3260 (2003).

    Google Scholar 

  41. Cai, L. et al. Effects of prolonged and burst swimming on subsequent burst swimming performance of Gymnocypris potanini firmispinatus (Actinopterygii, Cyprinidae). Hydrobiologia 843(1), 201–209 (2019).

    Google Scholar 

  42. Webb, P. W. Hydrodynamics and energetics of fish propulsion. Ottawa: Canada: Department of the Environment Fisheries and Marine Service.Bulletin 190 (1975).

  43. Wu, G., Yang, Y. & Zeng, L. Kinematics, hydrodynamics and energetic advantages of burst-and-coast swimming of koi carps (Cyprinus carpio koi). J. Exp. Biol. 210(Pt 12), 2181 (2007).

    Google Scholar 

  44. Blake, R. W. Fish Locomotion (Cambridge University Press, Cambridge, 1983).

    Google Scholar 

  45. Williams, J. G., Armstrong, G., Katopodis, C., Larinier, M. & Travade, F. Thinking like a fish: a key ingredient for development of effective fish passage facilities at river obstructions. River Res. Appl. 28(4), 407 (2012).

    Google Scholar 

  46. Wei, S., Suping, L., Gang, W., Yinxin, Z. & Xiaozhong, R. Research progress and prospect of fish swimming mechanism: a review. J. Dalian Ocean Univ. (3) 2024.

  47. Manduca, G. et al. A bioinspired control strategy ensures maneuverability and adaptability for dynamic environments in an underactuated robotic fish. J. Intell. Robot. Syst. 110(2), 69 (2024).

    Google Scholar 

  48. Chen, H., Li, W., Cui, W., Yang, P. & Chen, L. Multi-objective multidisciplinary design optimization of a robotic fish system. J. Mar. Sci. Eng. 9(5), 478 (2021).

    Google Scholar 

  49. Zhao, Q. et al. Fast-moving piezoelectric micro-robotic fish with double caudal fins. Robot. Auton. Syst. 140, 103 (2021).

    Google Scholar 

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Acknowledgements

This research was supported by the National Nature Science Foundation of China (Grant number: U2340218), the National Key R&D Program of China (Grant numbers: 2022YFE0117400, 2022YFC3203900).

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Contributions

Yiqun Hou and Feifei He wrote the main manuscript text, Xiang Wang, Xiaogang Wang ,Long Zhu and Lu Cai participated in the experiment and data processing. All authors reviewed the manuscript.

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Correspondence to
Feifei He.

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The authors declare no competing interests.

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The research was conducted in accordance with the Guide for the Care and Use of Laboratory Animals promulgated by the National Institutes of Health (NIH), 1996. All experimental protocols were approved by the Institute of Hydroecology, Ministry of Water Resources, and the Chinese Academy of Sciences (Approval number: 2023–05). Fishing during the study was approved by the local fishery administrations of the Agricultural and Rural Bureau of Wuhan Province. The sampling location is not privately owned or protected in any way and requires no specific permissions.

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Hou, Y., Wang, X., He, F. et al. Morphology and energetics of the wake behind a continuously swimming crucian carp at different flow velocities.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-46672-x

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  • DOI: https://doi.org/10.1038/s41598-026-46672-x

Keywords

  • Swimming
  • Flow field
  • Wake structure
  • Particle image velocimetry
  • Crucian carp


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