Reuse & Permissions

It is not necessary to obtain permission to reuse this article or its components as it is available under the terms of the Creative Commons Attribution 4.0 International license. This license permits unrestricted use, distribution, and reproduction in any medium, provided attribution to the author(s) and the published article's title, journal citation, and DOI are maintained. Please note that some figures may have been included with permission from other third parties. It is your responsibility to obtain the proper permission from the rights holder directly for these figures.

Export citation

Export citation

Choose format for download:

Download Citation
  • Open Access

Algal optics

Ming Yang*,†, Sumit Kumar Birwa*,‡, and Raymond E. Goldstein§

  • *These authors contributed equally to this work.
  • †Contact author: my365@cam.ac.uk
  • ‡Contact author: skb61@cam.ac.uk
  • §Contact author: R.E.Goldstein@damtp.cam.ac.uk

Phys. Rev. E 113, 024401 – Published 4 February, 2026

DOI: https://doi.org/10.1103/fp9l-zykg

Abstract

Nearly a decade ago, it was discovered that the spherical cell body of the alga Chlamydomonas reinhardtii can act as a lens to concentrate incoming light onto the cell's membrane-bound photoreceptor and thereby affect phototaxis. Since many nearly transparent cells in marine environments have complex, often nonaxisymmetric shapes, this observation raises fundamental, yet little-explored questions in biological optics about light refraction by the bodies of microorganisms. There are two distinct contexts for such questions: the absorption problem for incoming light, typified by photosynthetic activity taking place in the chloroplasts of green algae, and the emission problem for outgoing light, where the paradigm is bioluminescence emitted from scintillons within dinoflagellates. Here we examine both of these aspects of “algal optics” in the special case where the absorption or emission is localized in structures that are small relative to the overall organism size, taking into account both refraction and reflections at the cell-water boundary. Analytical and numerical results are developed for the distribution of light intensities inside and outside the body, and we establish certain duality relationships that connect the incoming and outgoing problems. For strongly nonspherical shapes, we find lensing effects that may have implications for photosynthetic activity and for the angular distribution of light emitted during bioluminescent flashes.

View figure in article

Physics Subject Headings (PhySH)

Article Text

References (62)

  1. J. Kessler, A. M. Nedelcu, C. A. Solari, and D. E. Shelton, Cells acting as lenses: A possible role for light in the evolution of morphological asymmetry in multicellular volvocine algae, evolutionary transitions to multicellular life, Adv. Marine Genomics 2, 225 (2015).
  2. N. Ueki, T. Ide, S. Mochiji, Y. Kobayashi, R. Tokutsu, N. Ohnishi, K. Yamaguchi, S. Shigenobu, K. Tanaka, J. Minagawa, T. Hisabori, M. Hirono, and K. Wakabayashi, Eyespot-dependent determination of the phototactic sign in Chlamydomonas reinhardtii, Proc. Natl. Acad. Sci. USA 113, 5299 (2016).
  3. A. Hallmann, Sensing a rainbow of colors: Algal photoreceptors, Front. Plant Sci. 16, 1684559 (2025).
  4. Aleksandra Szczerbiak, CC BY 4.0, via wikimedia commons, https://en.wikipedia.org/wiki/Pyrocystis_fusiformis.
  5. K. W. Foster and R. D. Smyth, Light antennas in phototactic algae, Microbiol. Rev. 44, 572 (1980).
  6. K. Schaller, R. David, and R. Uhl, How Chlamydomonas keeps track of the light once it has reached the right phototactic orientation, Biophys. J. 73, 1562 (1997).
  7. K. C. Leptos, M. Chioccioli, S. Furlan, A. I. Pesci, and R. E. Goldstein, Phototaxis of Chlamydomonas arises from a tuned adaptive photoresponse shared with multicellular Volvocine green algae, Phys. Rev. E 107, 014404 (2023).
  8. G. Jékely, Evolution of phototaxis, Phil. Trans. R. Soc. B 364, 2795 (2009).
  9. D. Francis, On the eyespot of the dinoflagellate, nematodinium, J. Expt. Biol. 47, 495 (1967).
  10. N. Schuergers, et al., Cyanobacteria use micro-optics to sense light direction, eLife 5, e12620 (2016).
  11. D. Nakane and T. Nishizaka, Asymmetric distribution of type IV pili triggered by directional light in unicellular cyanobacteria, Proc. Natl. Acad. Sci. USA 114, 6593 (2017).
  12. A. Wilde, et al., Light-controlled motility in prokaryotes and the problem of directional light perception, FEMS Microbiol. Rev. 41, 900 (2017).
  13. M. M. Ghobara, et al., On light and diatoms: A photonics and photobiology review, in Diatoms: Fundamentals and Applications, edited by J. Seckbach and R. Gordon (Wiley-Scrivener, Beverly, MA, 2019), pp. 129–189.
  14. E. De Tommasi, et al., Optics with diatoms: Towards efficient, bioinspired photonic devices at the micro-scale, optical methods for inspection, characterization, and imaging of biomaterials, SPIE 8792, 99 (2013).
  15. C. Maibohm, et al., Comparing optical properties of different species of diatoms, organic photonic materials and devices XVII, SPIE 9360, 93600B (2015).
  16. T. C. Vogelmann, Plant tissue optics, Annu. Rev. Plant Physiol. Plant Mol. Biol. 44, 231 (1993).
  17. V. R. Humphry, The effects of paraffin oil on phototropic and geotropic responses in Avena coleoptiles, Ann. Bot. 30, 39 (1966).
  18. D. S. Dennison, et al., The Phycomyces lens: Measurement of the sporangiophore intensity profile using a fiber optic microprobe, Planta 179, 1 (1989).
  19. Photon-vegetation Interactions: Applications in Optical Remote Sensing and Plant Ecology, edited by R. B. Myneni and J. Ross (Springer-Verlag, Berlin, 1991).
  20. R. A. Bone, et al., Epidermal cells functioning as lenses in leaves of tropical rain-forest shade plants, Appl. Opt. 24, 1408 (1985).
  21. C. R. Brodersen, et al., Do epidermal lens cells facilitate the absorptance of diffuse light? Amer. J. Botany 94, 1061 (2007).
  22. W. T. Plummer, J. G. Baker, and J. Van Tassell, Photographic optical systems with nonrotational aspheric surfaces, Appl. Opt. 38, 3572 (1999).
  23. O. Cakmakci, B. Moore, H. Foroosh, and J. P. Rolland, Optimal local shape description for rotationally non-symmetric optical surface design and analysis, Opt. Express 16, 1583 (2008).
  24. K. Fuerschbach, J. P. Rolland, and K. P. Thompson, A new family of optical systems employing φ-polynomial surfaces, Opt. Express 19, 21919 (2011).
  25. P. J. Smilie and T. J. Suleski, Variable-diameter refractive beam shaping with freeform optical surfaces, Opt. Lett. 36, 4170 (2011).
  26. E. Swift and W. R. Taylor, Bioluminescence and chloroplast movement in the dinoflagellate Pyrocystis lunula, J. Phycol. 3, 77 (1967).
  27. B. M. Sweeney, The circadian rhythms, bioluminescence, photosynthesis and organellar movements in the large dinoflagellate, Pyrocystis fusiformis, in International Cell Biology 1980–1981, edited by H. G. Schweiger (Springer, Berlin, 1981).
  28. K. Heimann, P. L. Klerks, and K. H. Hasenstein, Involvement of actin and microtubules in regulation of bioluminescence and translocation of chloroplasts in the dinoflagellate pyrocystis lunula, Bot. Mar. 52, 170 (2009).
  29. F. C. Stephens, Variability of spectral absorption efficiency within living cells of pyrocystis lunula (dinophyta), Mar. Biol. 122, 325 (1995).
  30. S. H. D. Haddock, M. A. Moline, and J. F. Case, Bioluminescence in the sea, Annu. Rev. Mar. Sci. 2, 443 (2010).
  31. M. I. Latz, M. Bovard, V. VanDelinder, E. Segre, J. Rohr, and A. Groisman, Bioluminescent response of individual dinoflagellate cells to hydrodynamic stress measured with millisecond resolution in a microfluidic device, J. Expt. Biol. 211, 2865 (2008).
  32. M. Jalaal, N. Schramma, A. Dode, H. de Maleprade, C. Raufaste, and R. E. Goldstein, Stress-induced dinoflagellate bioluminescence at the single cell level, Phys. Rev. Lett. 125, 028102 (2020).
  33. J. S. Font-Muñoz, J. Arrieta, M. Sourisseau, I. Tuval, and G. Basterretxea, Anisotropic fluorescence by diatoms modifies the underwater oceanic light field, bioRxiv (2025).
  34. Y.-T. Tu, et al., Ray optics simulation, Zenodo (2016), https://zenodo.org/records/17718563.
  35. X. Quan and E. S. Fry, Empirical equation for the index of refraction of seawater, Appl. Opt. 34, 3477 (1995).
  36. J. D. Jackson, Classical Electrodynamics, 3rd ed. (Wiley, New York, 1999).
  37. S. K. Birwa, M. Yang, A. I. Pesci, and R. E. Goldstein, Theory of cell body lensing and phototaxis sign reversal in “eyeless” mutants of chlamydomonas (unpublished).
  38. R. E. Goldstein, Green algae as model organisms for biological fluid dynamics, Annu. Rev. Fluid Mech. 47, 343 (2015).
  39. J. Chaves, Introduction to Nonimaging Optics, 2nd ed. (CRC Press, Boca Raton, FL, 2016).
  40. M. F. Modest, Radiative Heat Transfer, 3rd ed. (Academic Press, Oxford, UK, 2013).
  41. E. H. Harris, The Chlamydomonas Sourcebook (Academic Press, Oxford, UK, 2009).
  42. D. L. Kirk, Volvox: Molecular-genetic Origins of Multicellularity and Cellular Differentiation (Cambridge University Press, Cambridge, UK, 1998).
  43. C. J. M. Hoppe, et al., Photosynthetic light requirement near the theoretical minimum detected in arctic microalgae, Nat. Commun. 15, 7385 (2024).
  44. M. Poirier, P. Osmers, K. Wilkins, R. M. Morgan-Kiss, and M. Cvetkovska, Aberrant light sensing and motility in the green alga chlamydomonas priscuii from the ice-covered Antarctic lake bonney, Plant Sig. Behav. 18, 2184588 (2023).
  45. Photobiology: The Science of Life and Light, 2nd ed., edited by L. O. Björn (Springer, New York, 2008).
  46. M. Wada, et al., Chloroplast movement, Annu. Rev. Plant Biol. 54, 455 (2003).
  47. N. Schramma, C. P. Israëls, and M. Jalaal, Chloroplasts in plant cells show active glassy behavior under low-light conditions, Proc. Natl. Acad. Sci. USA 120, e2216497120 (2023).
  48. N. Schramma, G. C. Canales, and M. Jalaal, Light-regulated chloroplast morphodynamics in a single-celled dinoflagellate, Proc. Natl. Acad. Sci. USA 121, e2411725121 (2024).
  49. L. Karp-Boss, E. Boss, and P. A. Jumars, Motion of dinoflagellates in a simple shear flow, Limn. Ocean. 45, 1594 (2000).
  50. J. S. Guasto, R. Rusconi, and R. Stocker, Fluid mechanics of planktonic microorganisms, Annu. Rev. Fluid Mech. 44, 373 (2012).
  51. J. S. Font-Muñoz, M. Sourisseau, A. Cohen-Sánchez, I. Tuval, and G. Basterretxea, Pelagic diatoms communicate through synchronized beacon natural fluorescence signaling, Sci. Adv. 7, eabj5230 (2021).
  52. J. S. Font-Muñoz, M. Jaubert, M. Sourisseau, I. Tuval, B. Bailleul, C. Duchêne, G. Basterretxea, and A. Falciatore, Phytochromes enable social behavior in marine diatoms, bioRxiv (2024).
  53. I. Eisenmann, A. L'Homme, A. Lehlou, S. Bujaldon, T. Le Saux, B. Bailleul, N. Desprat, and R. Jeanneret, Light-induced phase separation and finite wavelength selection in photophobic microalgae, Phys. Rev. Lett. 135, 148401 (2025).
  54. I. Eisenmann, M. Vona, N. Desprat, T. Ishikawa, E. Lauga, and R. Jeanneret, Pure hydrodynamic instabilities in active jets of puller microalgae, Phys. Rev. Lett. 135, 198301 (2025).
  55. R. D. Airan, K. R. Thompson, L. E. Fenno, H. Bernstein, and K. Deisseroth, Temporally precise in vivo control of intracellular, Nature (London) 458, 1025 (2009).
  56. J. Arrieta, A. Barreira, M. Chioccioli, M. Polin, and I. Tuval, Phototaxis beyond turning: Persistent accumulation and response acclimation of the microalga Chlamydomonas reinhardtii, Sci. Rep. 7, 3447 (2017).
  57. T. Laroussi, M. Jarrahi, and G. Amselem, Short-term memory effects in the phototactic behavior of microalgae, Soft Matter 20, 3996 (2024).
  58. Z. Wang and A. C. H. Tsang, Intermediate light adaptation induces oscillatory phototaxis switching and pattern formation in Chlamydomonas, Proc. Natl. Acad. Sci. USA 122, e2425369122 (2025).
  59. J. Morelle, A. Bastos, S. Frankenbach, J. C. Frommlet, D. A. Campbell, J. Lavaud, and J. Serôdio, The photoprotective behavior of a motile benthic diatom as elucidated from the interplay between cell motility and physiological responses to a light microgradient using a novel experimental setup, Micro. Ecol. 87, 40 (2024).
  60. S. Johnsen, The Optics of Life: A Biologist's Guide to Light in Nature (Princeton University Press, Princeton, NJ, 2012).
  61. C. Lozano, B. ten Hagen, H. Löwen, and C. Bechinger, Phototaxis of synthetic microswimmers in optical landscapes, Nat. Commun. 7, 12828 (2016).
  62. M. Yang, S. K. Birwa, and R. E. Goldstein, Data for “Algal optics,” Zenodo (2025), https://zenodo.org/records/15552607.

Outline

Information

Sign In to Your Journals Account

Filter

Filter

Article Lookup

Enter a citation