Experimental investigation of turbulent flow over live mussels


Unionids have been described as ecosystem engineers capable of affecting the local food web as well as the hydrodynamics, passively and actively. In this study, we perform particle image velocimetry measurements to characterize the flow around a live Amblema plicata using natural sediments as tracer particles under two specific flow rates and orientations i.e., facing upstream and downstream, to understand the interaction between the organism and flow surrounding it. The behavior of the mussel is also quantified under all the above experimental conditions using a hall-effect gape sensor that captures valve motion. The flow measurements show clear variations between the two orientations tested, with the presence of a low velocity region behind the mussel extending beyond the tested field of view for the upstream orientation. In contrast, the flow in the wake of the downstream orientation recovers very quickly to the incoming flow strength. The inspection of the vorticity contours around the mussel also highlights clear differences between orientations, with the structure of a shear layer under the upstream orientation changing to a region of concentrated vorticity near the siphons for the downstream cases. The variation of the net flux through the mussel along with the frequency of partial closures captured by the gape sensor measurements illustrate a potential added bioenergetic cost to the mussel feeding when facing downstream, especially at higher flow speeds.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7



Significance level in Tukey’s post-hoc test

h :

Exposed height of mussel above the sediment bed

Ω :

Mean vorticity

\(\phi = \frac{1}{L}\mathop {\oint }\limits_{L}^{{}} \vec{u} \cdot \hat{n} dl\) :

Average net flux of water through the mussel

L :

Length along the shell of the mussel

\(\hat{n}\) :

Normal vector around the mussel shell

\(\vec{u}\) :

Velocity vector in two dimensions (x and z directions)

U :

Mean streamwise velocity (x-direction)

U :

Free stream velocity

x :

Dimension in the streamwise direction

z :

Dimension in the wall normal direction


  1. 1.

    Jones CG, Lawton JH, Shachak M (1994) Organisms as ecosystem engineers. In: Samson FB, Knopf FL (eds) Ecosystem management. Springer, New York, pp 130–147. https://doi.org/10.1007/978-1-4612-4018-1_14

    Google Scholar 

  2. 2.

    Butler DR, Sawyer CF (2012) Introduction to the special issue: zoogeomorphology and ecosystem engineering. Geomorphology 157–158:1–5. https://doi.org/10.1016/j.geomorph.2012.02.027

    Article  Google Scholar 

  3. 3.

    Polvi LE, Sarneel JM (2017) Ecosystem engineers in rivers: an introduction to how and where organisms create positive biogeomorphic feedbacks. Wiley Interdiscip Rev Water 5:e1271. https://doi.org/10.1002/wat2.1271

    Article  Google Scholar 

  4. 4.

    Spooner DE, Vaughn CC (2008) A trait-based approach to species roles in stream ecosystems: climate change, community structure, and material cycling. Oecologia 158:307–317. https://doi.org/10.1007/s00442-008-1132-9

    Article  Google Scholar 

  5. 5.

    Newton TJ, Woolnough DA, Strayer DL (2008) Using landscape ecology to understand and manage freshwater mussel populations. J N Am Benthol Soc 27:424–439. https://doi.org/10.1899/07-076.1

    Article  Google Scholar 

  6. 6.

    Haag WR (2012) North American freshwater mussels: natural history, ecology, and conservation. Cambridge University Press, New York

    Google Scholar 

  7. 7.

    Hardison BS, Layzer JB (2001) Relations between complex hydraulics and the localized distribution of mussels in three regulated rivers. Regul Rivers Res Manag 17:77–84. https://doi.org/10.1002/1099-1646(200101/02)17:1%3c77:AID-RRR604%3e3.3.CO;2-J

    Article  Google Scholar 

  8. 8.

    Zigler SJ, Newton TJ, Steuer JJ, Bartsch MR, Sauer JS (2008) Importance of physical and hydraulic characteristics to unionid mussels: a retrospective analysis in a reach of large river. Hydrobiologia 598:343–360. https://doi.org/10.1007/s10750-007-9167-1

    Article  Google Scholar 

  9. 9.

    Hornbach DJ, Kurth VJ, Hove MC (2010) Variation in freshwater mussel shell sculpture and shape along a river gradient. Am Midl Nat 164:22–36. https://doi.org/10.1674/0003-0031-164.1.22

    Article  Google Scholar 

  10. 10.

    Allen DC, Vaughn CC (2010) Complex hydraulic and substrate variables limit freshwater mussel species richness and abundance. J N Am Benthol Soc 29:383–394. https://doi.org/10.1899/09-024.1

    Article  Google Scholar 

  11. 11.

    Hart DD, Finelli CM (1999) Physical-biological coupling in streams: the pervasive effects of flow on benthic organisms. Annu Rev Ecol Syst 30:363–395. https://doi.org/10.1146/annurev.ecolsys.30.1.363

    Article  Google Scholar 

  12. 12.

    Statzner B (2008) How views about flow adaptations of benthic stream invertebrates changed over the last century. Int Rev Hydrobiol 93:593–605. https://doi.org/10.1002/iroh.200711018

    Article  Google Scholar 

  13. 13.

    Rice SP, Lancaster J, Kemp P (2010) Experimentation at the interface of fluvial geomorphology, stream ecology and hydraulic engineering and the development of an effective, interdisciplinary river science. Earth Surf Process Landf 35:64–77. https://doi.org/10.1002/esp

    Article  Google Scholar 

  14. 14.

    Trinci G, Harvey GL, Henshaw AJ, Bertoldi W, Hölker F (2017) Life in turbulent flows: interactions between hydrodynamics and aquatic organisms in rivers. Wiley Interdiscip Rev Water 4:e1213. https://doi.org/10.1002/wat2.1213

    Article  Google Scholar 

  15. 15.

    Baker SM, Hornbach DJ (2000) Physiological status and biochemical composition of a natural population of unionid mussels (Amblema plicata) infested by zebra mussels (Dreissena polymorpha). Am Midl Nat 143:443–452. https://doi.org/10.1674/0003-0031(2000)143[0443:PSABCO]2.0.CO;2

    Article  Google Scholar 

  16. 16.

    Baker SM, Hornbach DJ (2008) Zebra mussels (Dreissena polymorpha) attached to native mussels (Unionidae) or inanimate substrates: comparison of physiological rates and biochemical composition. Am Midl Nat 160:20–28. https://doi.org/10.1674/0003-0031(2008)160[20:ZMDPAT]2.0.CO;2  

    Article  Google Scholar 

  17. 17.

    Sansom BJ, Atkinson JF, Bennett SJ (2018) Modulation of near-bed hydrodynamics by freshwater mussels in an experimental channel. Hydrobiologia 810:449–463. https://doi.org/10.1007/s10750-017-3172-9

    Article  Google Scholar 

  18. 18.

    Albertson LK, Allen DC, Trexler JC (2015) Meta-analysis: abundance, behavior, and hydraulic energy shape biotic effects on sediment transport in streams. Ecology 96:1329–1339. https://doi.org/10.1890/13-2138.1

    Article  Google Scholar 

  19. 19.

    Nishizaki M, Ackerman JD (2017) Mussels blow rings: jet behavior affects local mixing. Limnol Oceanogr 62:125–136. https://doi.org/10.1002/lno.10380

    Article  Google Scholar 

  20. 20.

    Jones J, Ahlstedt S, Ostby B, Beaty B, Pinder M, Eckert N, Butler R, Hubbs D, Walker C, Hanlon S, Schmerfeld J, Neves R (2014) Clinch river freshwater mussels upstream of Norris reservoir, Tennessee and Virginia: a quantitative assessment from 2004 to 2009. J Am Water Resour Assoc 50:820–836. https://doi.org/10.1111/jawr.12222

    Article  Google Scholar 

  21. 21.

    Daniel WM, Brown KM (2014) The role of life history and behavior in explaining unionid mussel distributions. Hydrobiologia 734:57–68. https://doi.org/10.1007/s10750-014-1868-7

    Article  Google Scholar 

  22. 22.

    Atkinson CL, Vaughn CC, Forshay KJ, Cooper JT (2013) Aggregated filter-feeding consumers alter nutrient limitation: consequences for ecosystem and community dynamics. Ecology 94:1359–1369. https://doi.org/10.1890/12-1531.1

    Article  Google Scholar 

  23. 23.

    Strayer D (2014) Understanding how nutrient cycles and freshwater mussels (unionida) affect one another. Hydrobiologia 735:277–292. https://doi.org/10.1007/s10750-013-1461-5

    Article  Google Scholar 

  24. 24.

    Hoellein TJ, Zarnoch CB, Bruesewitz DA, DeMartini J (2017) Contributions of freshwater mussels (Unionidae) to nutrient cycling in an urban river: filtration, recycling, storage, and removal. Biogeochemistry 135:307–324. https://doi.org/10.1007/s10533-017-0376-z

    Article  Google Scholar 

  25. 25.

    Atkinson CL, Sansom BJ, Vaughn CC, Forshay KJ (2018) Consumer aggregations drive nutrient dynamics and ecosystem metabolism in nutrient-limited systems. Ecosystems 21:521–535. https://doi.org/10.1007/s10021-017-0166-4

    Article  Google Scholar 

  26. 26.

    Trentman MT, Atkinson CL, Brant JD (2018) Native freshwater mussel effects on nitrogen cycling: impacts of nutrient limitation and biomass dependency. Freshw Sci 37:276–286. https://doi.org/10.1086/697293

    Article  Google Scholar 

  27. 27.

    Atkinson CL, Kelly JF, Vaughn CC (2014) Tracing consumer-derived nitrogen in riverine food webs. Ecosystems 17:485–496. https://doi.org/10.1007/s10021-013-9736-2

    Article  Google Scholar 

  28. 28.

    Vaughn CC, Gido KB, Spooner DE (2004) Ecosystem processes performed by unionid mussels in stream mesocosms: species roles and effects of abundance. Hydrobiologia 527:35–47. https://doi.org/10.1023/B:HYDR.0000043180.30420.00

    Article  Google Scholar 

  29. 29.

    Tevesz MJS, Cornelius DW, Fisher JB (1985) Life habits and distribution of riverine Lampsilis radiata luteola (Mollusca: bivalvia). Kirtlandia 41:27–34

    Google Scholar 

  30. 30.

    Di Maio J, Corkum LD (1997) Patterns of orientation in unionids as a function of rivers with differing hydrological variability. J Molluscan Stud 63:531–539. https://doi.org/10.1093/mollus/63.4.531

    Article  Google Scholar 

  31. 31.

    Perles SJ, Christian AD, Berg DJ (2003) Vertical migration, orientation, aggregation, and fecundity of the freshwater mussel Lampsilis siliquoidea. Ohio J Sci 103(4):73–78

    Google Scholar 

  32. 32.

    Constantinescu G, Miyawaki S, Liao Q (2012) Flow and turbulence structure past a cluster of freshwater mussels. J Hydraul Eng 139:347–358. https://doi.org/10.1061/(ASCE)HY.1943-7900.0000692

    Article  Google Scholar 

  33. 33.

    Riisgård HU, Jørgensen BH, Lundgreen K, Storti F, Walther JH, Meyer KE, Larsen PS (2011) The exhalent jet of mussels mytilus edulis. Mar Ecol Prog Ser 437:147–164. https://doi.org/10.3354/meps09268

    Article  Google Scholar 

  34. 34.

    Frank DM, Ward JE (2008) Application of particle image velocimetry to the study of suspension feeding in marine invertebrates. Mar Freshw Behav Physiol 41:1–18. https://doi.org/10.1080/10236240801896207

    Article  Google Scholar 

  35. 35.

    Morales Y, Weber LJ, Mynett AE, Newton TJ (2006) Mussel dynamics model: a hydroinformatics tool for analyzing the effects of different stressors on the dynamics of freshwater mussel communities. Ecol Modell 197:448–460. https://doi.org/10.1016/j.ecolmodel.2006.03.018

    Article  Google Scholar 

  36. 36.

    Yunus AC, Cimbala JM (2006) Fluid mechanics: fundamentals and applications, 3rd edn. McGraw-Hill Education, Boston

    Google Scholar 

  37. 37.

    Hartmann JT, Beggel S, Auerswald K, Geist J (2016) Determination of the most suitable adhesive for tagging freshwater mussels and its use in an experimental study of filtration behaviour and biological rhythm. J Molluscan Stud 82:415–421. https://doi.org/10.1093/mollus/eyw003

    Article  Google Scholar 

  38. 38.

    Robson AA, Thomas GR, De Leaniz CG, Wilson RP (2009) Valve gape and exhalant pumping in bivalves: optimization of measurement. Aquat Biol 6:191–200. https://doi.org/10.3354/ab00128

    Article  Google Scholar 

  39. 39.

    Wilson R, Reuter P, Wahl M (2005) Muscling in on mussels: new insights into bivalve behaviour using vertebrate remote-sensing technology. Mar Biol 147:1165–1172. https://doi.org/10.1007/s00227-005-0021-6

    Article  Google Scholar 

  40. 40.

    Raffel M, Willert CE, Scarano F, Kähler CJ, Wereley ST, Kompenhans J (2018) Particle image velocimetry: a practical guide. Springer, Berlin

    Google Scholar 

  41. 41.

    Roth GI, Katz J (2001) Five techniques for increasing the speed and accuracy of PIV interrogation. Meas Sci Technol 12:238–245. https://doi.org/10.1088/0957-0233/12/3/302

    Article  Google Scholar 

  42. 42.

    Hasler CT, Hannan KD, Jeffrey JD, Suski CD (2017) Valve movement of three species of north American freshwater mussels exposed to elevated carbon dioxide. Environ Sci Pollut Res 24:15567–15575. https://doi.org/10.1007/s11356-017-9160-9

    Article  Google Scholar 

Download references


Funding for this project was provided by the Minnesota Environment and Natural Resources Trust Fund as recommended by the Legislative-Citizen Commission on Minnesota Resources (LCCMR). Gape sensors were built by Chris Ellis, with advice from Joel Allen and Anton Kruger. Mark Hove assisted with mussel collection and housing. SAFL Technical Staff including Eric Steen, Dick Christopher and Ben Erickson along with undergraduate research assistants assisted with flume setup. The authors would also like to thank Brandon Samson for some fruitful discussions.

Author information



Corresponding author

Correspondence to Jiarong Hong.

Additional information

Publisher's Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary material 1 (DOCX 322 kb)

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Kumar, S.S., Kozarek, J., Hornbach, D. et al. Experimental investigation of turbulent flow over live mussels. Environ Fluid Mech 19, 1417–1430 (2019). https://doi.org/10.1007/s10652-019-09664-2

Download citation


  • Freshwater mussel
  • PIV measurements
  • Gape sensors
  • Turbulent flows
  • Hydrodynamic interactions