Abstract

Dramatic and rapid changes in the intensity and spectrum of light under water at dusk and dawn are well known, but reports regarding the light’s polarization at these periods are sparse. Using a rapid spectropolarimeter, we examined the spatial and spectral characteristics of the underwater polarization patterns from sunrise to midday and compared them with a Rayleigh-based model for e-vector orientation and percent polarization. With the Sun near the horizon, the underwater polarization patterns were distinctive. Unlike the polarization at small solar zenith angles, the underwater polarization at large solar zenith angles cannot be predicted by simple Rayleigh scattering, most likely because of the relatively high contribution of skylight. At sunrise, the underwater polarization pattern outside of Snell’s window differed from that found during the day in percent polarization, spatial distribution, and wavelength dependence. These unique polarization characteristics may provide a polarization-sensitive animal with a distinct cue for mediating dial vertical migration performed by plankton or with another timing signal.

© 2007 Optical Society of America

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  1. W. N. McFarland, "Light in the sea—correlations with behaviors of fishes and invertebrates," Am. Zool. 26, 389-401 (1986).
  2. W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.
  3. T. H. Waterman, "Polarization sensitivity," in Comparative Physiology and Evolution of Vision in Invertebrates. B: Invertebrate Visual Centers and Behavior I, H.Autrum, ed. (Springer-Verlag, 1981), pp. 281-469.
  4. I. Novales Flamarique and C. W. Hawryshyn, "Is the use of underwater polarized light by fish restricted to crepuscular time periods?" Vision Res. 37, 975-989 (1997).
    [CrossRef] [PubMed]
  5. T. H. Waterman and W. E. Westell, "Quantitative effect of the sun's position on submarine light polarization," J. Mar. Res. 15, 149-169 (1956).
  6. W. G. Egan, Photometry and Polarization in Remote Sensing (Elsevier, 1985).
  7. A. Ivanoff and T. H. Waterman, "Elliptical polarization of submarine illumination," J. Mar. Res. 16, 255-282 (1958).
  8. G. Horvath and D. Varjú, Polarized Light in Animal Vision: Polarization Patterns in Nature (Springer-Verlag, 2004).
  9. V. A. Timofeyeva, "Plane of vibrations of polarized light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 5, 1049-1057 (1969).
  10. T. H. Waterman, "Polarization patterns in submarine illumination," Science 120, 927-932 (1954).
    [CrossRef] [PubMed]
  11. T. W. Cronin and N. Shashar, "The linearly polarized light field in clear, tropical marine waters: spatial and temporal variation of light intensity, degree of polarization and e-vector angle," J. Exp. Biol. 204, 2461-2467 (2001).
    [PubMed]
  12. A. Ivanoff and T. H. Waterman, "Factors, mainly depth and wavelength, affecting the degree of underwater light polarization," J. Mar. Res. 16, 283-307 (1958).
  13. V. A. Timofeyeva, "The degree of polarization of light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 6, 513-522 (1970).
  14. G. Horváth and D. Varjú, "Underwater refraction-polarization patterns of skylight perceived by aquatic animals through Snell's window of the flat water surface," Vision Res. 35, 1651-1666 (1995).
    [CrossRef] [PubMed]
  15. S. Sabbah, A. Barta, J. Gal, G. Horváth, and N. Shashar, "Experimental and theoretical study of skylight polarization transmitted through Snell's window of a flat water surface," J. Opt. Soc. Am. A 23, 1978-1988 (2006).
    [CrossRef]
  16. J. T. Adams, E. Aas, N. K. Hojerslev, and B. Lundgren, "Comparison of radiance and polarization values observedin the Mediterranean Sea and simulated in a Monte Carlo model," Appl. Opt. 41, 2724-2733 (2002).
    [CrossRef] [PubMed]
  17. G. W. Kattawar, G. N. Plass, and J. J. A. Guinn, "Monte Carlo calculations of the polarization of radiation in the Earth's atmosphere-ocean system," J. Phys. Oceanogr. 3, 353-372 (1973).
    [CrossRef]
  18. J. T. Adams and G. W. Kattawar, "Neutral points in an atmosphere-ocean system. 1: Upwelling light field," Appl. Opt. 36, 1976-1986 (1997).
    [CrossRef] [PubMed]
  19. R. Wehner, "Polarization vision—a uniform sensory capacity?" J. Exp. Biol. 204, 2589-2596 (2001).
    [PubMed]
  20. S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.
  21. C. Groot, "On the orientation of young sockeye salmon (Oncorhynchus nerka), during their seaward migration out of lakes," Behaviour 14, 198 (1965).
  22. R. B. Forward, K. W. Horch, and T. H. Waterman, "Visual orientation at the water surface by the teleost Zenarchopterus," Biol. Bull. 143, 112-126 (1972).
    [CrossRef]
  23. R. B. Forward and T. H. Waterman, "Evidence for e-vector and light intensity pattern discrimination by the teleost Demogenys," J. Comp. Physiol., A 87, 189-202 (1973).
    [CrossRef]
  24. H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
    [CrossRef]
  25. C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
    [CrossRef]
  26. C. W. Hawryshyn, "Polarization vision in fish," Am. Sci. 80, 479-491 (1992).
  27. R. Schwind, "Daphnia pulex swims towards the most strongly polarized light—a response that leads to 'shore flight'," J. Exp. Biol. 202, 3631-3635 (1999).
    [PubMed]
  28. D. A. Ritz, "Polarized-light responses in the shrimp Palaemonetes vulgari (Say)," J. Exp. Mar. Biol. Ecol. 154, 245-250 (1991).
    [CrossRef]
  29. S. M. Goddard and R. B. Forward, "The role of the underwater polarized light pattern, in sun compass navigation of the grass shrimp, Palaemonetes vulgaris," J. Comp. Physiol., A 169, 479-491 (1991).
    [CrossRef]
  30. E. J. Denton and J. A. C. Nicol, "Polarization of light reflected from the silvery exterior of the bleak Alburnus alburnus," J. Mar. Biol. Assoc. U.K. 45, 705-709 (1965).
    [CrossRef]
  31. N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
    [CrossRef] [PubMed]
  32. N. Shashar, R. T. Hanlon, and A. D. Petz, "Polarization vision helps detect transparent prey," Nature 393, 222-223 (1998).
    [CrossRef]
  33. I. Novales Flamarique and H. I. Browman, "Foraging and prey-search behaviour of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).
  34. B. L. Umminger, "Polarotaxis in copepods. I. An endogenous rhythm in polarotaxis in Cyclops vernalis and its relation to vertical migration," Biol. Bull. 135, 239-251 (1968).
    [CrossRef]
  35. N. Shashar, S. Sabbah, and T. W. Cronin, "Transmission of linearly polarized light in sea water implications for polarization signaling," J. Exp. Biol. 207, 3619-3628 (2004).
    [CrossRef] [PubMed]
  36. S. Sabbah and N. Shashar, "Underwater light polarization and radiance fluctuations induced by surface waves," Appl. Opt. 45, 4726-4739 (2006).
    [CrossRef] [PubMed]
  37. L. B. Wolff and A. G. Andreou, "Polarization camera sensors," Image Vis. Comput. 13, 497-509 (1995).
    [CrossRef]
  38. C. D. Mobley, Light and Water (Academic, 1994).
  39. E. Boss, School of Marine Sciences, University of Maine, Orono, Maine, 04469 (personal communication, 2005).
  40. A. Ivanoff, "Polarization measurements in the sea," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 151-175.
  41. V. A. Timofeyeva, "Optics of turbid waters (results of laboratory studies)," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 177-219.
  42. E. S. Hobson, "Trophic relationships of fishes specialized to feed on zooplankters above coral reefs," in The Ecology of Fishes on Coral Reefs (Academic, 1991), pp. 69-94.

2006

2005

S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.

E. Boss, School of Marine Sciences, University of Maine, Orono, Maine, 04469 (personal communication, 2005).

2004

G. Horvath and D. Varjú, Polarized Light in Animal Vision: Polarization Patterns in Nature (Springer-Verlag, 2004).

N. Shashar, S. Sabbah, and T. W. Cronin, "Transmission of linearly polarized light in sea water implications for polarization signaling," J. Exp. Biol. 207, 3619-3628 (2004).
[CrossRef] [PubMed]

2002

2001

T. W. Cronin and N. Shashar, "The linearly polarized light field in clear, tropical marine waters: spatial and temporal variation of light intensity, degree of polarization and e-vector angle," J. Exp. Biol. 204, 2461-2467 (2001).
[PubMed]

R. Wehner, "Polarization vision—a uniform sensory capacity?" J. Exp. Biol. 204, 2589-2596 (2001).
[PubMed]

I. Novales Flamarique and H. I. Browman, "Foraging and prey-search behaviour of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).

2000

N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
[CrossRef] [PubMed]

1999

R. Schwind, "Daphnia pulex swims towards the most strongly polarized light—a response that leads to 'shore flight'," J. Exp. Biol. 202, 3631-3635 (1999).
[PubMed]

W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.

1998

N. Shashar, R. T. Hanlon, and A. D. Petz, "Polarization vision helps detect transparent prey," Nature 393, 222-223 (1998).
[CrossRef]

1997

I. Novales Flamarique and C. W. Hawryshyn, "Is the use of underwater polarized light by fish restricted to crepuscular time periods?" Vision Res. 37, 975-989 (1997).
[CrossRef] [PubMed]

J. T. Adams and G. W. Kattawar, "Neutral points in an atmosphere-ocean system. 1: Upwelling light field," Appl. Opt. 36, 1976-1986 (1997).
[CrossRef] [PubMed]

1995

G. Horváth and D. Varjú, "Underwater refraction-polarization patterns of skylight perceived by aquatic animals through Snell's window of the flat water surface," Vision Res. 35, 1651-1666 (1995).
[CrossRef] [PubMed]

L. B. Wolff and A. G. Andreou, "Polarization camera sensors," Image Vis. Comput. 13, 497-509 (1995).
[CrossRef]

1994

C. D. Mobley, Light and Water (Academic, 1994).

1992

C. W. Hawryshyn, "Polarization vision in fish," Am. Sci. 80, 479-491 (1992).

1991

D. A. Ritz, "Polarized-light responses in the shrimp Palaemonetes vulgari (Say)," J. Exp. Mar. Biol. Ecol. 154, 245-250 (1991).
[CrossRef]

S. M. Goddard and R. B. Forward, "The role of the underwater polarized light pattern, in sun compass navigation of the grass shrimp, Palaemonetes vulgaris," J. Comp. Physiol., A 169, 479-491 (1991).
[CrossRef]

E. S. Hobson, "Trophic relationships of fishes specialized to feed on zooplankters above coral reefs," in The Ecology of Fishes on Coral Reefs (Academic, 1991), pp. 69-94.

1990

C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
[CrossRef]

1986

W. N. McFarland, "Light in the sea—correlations with behaviors of fishes and invertebrates," Am. Zool. 26, 389-401 (1986).

1985

W. G. Egan, Photometry and Polarization in Remote Sensing (Elsevier, 1985).

1981

T. H. Waterman, "Polarization sensitivity," in Comparative Physiology and Evolution of Vision in Invertebrates. B: Invertebrate Visual Centers and Behavior I, H.Autrum, ed. (Springer-Verlag, 1981), pp. 281-469.

1974

A. Ivanoff, "Polarization measurements in the sea," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 151-175.

V. A. Timofeyeva, "Optics of turbid waters (results of laboratory studies)," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 177-219.

1973

R. B. Forward and T. H. Waterman, "Evidence for e-vector and light intensity pattern discrimination by the teleost Demogenys," J. Comp. Physiol., A 87, 189-202 (1973).
[CrossRef]

H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
[CrossRef]

G. W. Kattawar, G. N. Plass, and J. J. A. Guinn, "Monte Carlo calculations of the polarization of radiation in the Earth's atmosphere-ocean system," J. Phys. Oceanogr. 3, 353-372 (1973).
[CrossRef]

1972

R. B. Forward, K. W. Horch, and T. H. Waterman, "Visual orientation at the water surface by the teleost Zenarchopterus," Biol. Bull. 143, 112-126 (1972).
[CrossRef]

1970

V. A. Timofeyeva, "The degree of polarization of light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 6, 513-522 (1970).

1969

V. A. Timofeyeva, "Plane of vibrations of polarized light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 5, 1049-1057 (1969).

1968

B. L. Umminger, "Polarotaxis in copepods. I. An endogenous rhythm in polarotaxis in Cyclops vernalis and its relation to vertical migration," Biol. Bull. 135, 239-251 (1968).
[CrossRef]

1965

E. J. Denton and J. A. C. Nicol, "Polarization of light reflected from the silvery exterior of the bleak Alburnus alburnus," J. Mar. Biol. Assoc. U.K. 45, 705-709 (1965).
[CrossRef]

C. Groot, "On the orientation of young sockeye salmon (Oncorhynchus nerka), during their seaward migration out of lakes," Behaviour 14, 198 (1965).

1958

A. Ivanoff and T. H. Waterman, "Factors, mainly depth and wavelength, affecting the degree of underwater light polarization," J. Mar. Res. 16, 283-307 (1958).

A. Ivanoff and T. H. Waterman, "Elliptical polarization of submarine illumination," J. Mar. Res. 16, 255-282 (1958).

1956

T. H. Waterman and W. E. Westell, "Quantitative effect of the sun's position on submarine light polarization," J. Mar. Res. 15, 149-169 (1956).

1954

T. H. Waterman, "Polarization patterns in submarine illumination," Science 120, 927-932 (1954).
[CrossRef] [PubMed]

Aas, E.

Adams, J. T.

Andreou, A. G.

L. B. Wolff and A. G. Andreou, "Polarization camera sensors," Image Vis. Comput. 13, 497-509 (1995).
[CrossRef]

Arnold, M. G.

C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
[CrossRef]

Barta, A.

Boal, J. G.

N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
[CrossRef] [PubMed]

Boss, E.

E. Boss, School of Marine Sciences, University of Maine, Orono, Maine, 04469 (personal communication, 2005).

Bowering, D.

C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
[CrossRef]

Browman, H. I.

I. Novales Flamarique and H. I. Browman, "Foraging and prey-search behaviour of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).

Cole, R. L.

C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
[CrossRef]

Cronin, T. W.

N. Shashar, S. Sabbah, and T. W. Cronin, "Transmission of linearly polarized light in sea water implications for polarization signaling," J. Exp. Biol. 207, 3619-3628 (2004).
[CrossRef] [PubMed]

T. W. Cronin and N. Shashar, "The linearly polarized light field in clear, tropical marine waters: spatial and temporal variation of light intensity, degree of polarization and e-vector angle," J. Exp. Biol. 204, 2461-2467 (2001).
[PubMed]

Denton, E. J.

E. J. Denton and J. A. C. Nicol, "Polarization of light reflected from the silvery exterior of the bleak Alburnus alburnus," J. Mar. Biol. Assoc. U.K. 45, 705-709 (1965).
[CrossRef]

Egan, W. G.

W. G. Egan, Photometry and Polarization in Remote Sensing (Elsevier, 1985).

Erlick, C.

S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.

Forward, R. B.

S. M. Goddard and R. B. Forward, "The role of the underwater polarized light pattern, in sun compass navigation of the grass shrimp, Palaemonetes vulgaris," J. Comp. Physiol., A 169, 479-491 (1991).
[CrossRef]

R. B. Forward and T. H. Waterman, "Evidence for e-vector and light intensity pattern discrimination by the teleost Demogenys," J. Comp. Physiol., A 87, 189-202 (1973).
[CrossRef]

R. B. Forward, K. W. Horch, and T. H. Waterman, "Visual orientation at the water surface by the teleost Zenarchopterus," Biol. Bull. 143, 112-126 (1972).
[CrossRef]

Gal, J.

Gensler, P.

H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
[CrossRef]

Goddard, S. M.

S. M. Goddard and R. B. Forward, "The role of the underwater polarized light pattern, in sun compass navigation of the grass shrimp, Palaemonetes vulgaris," J. Comp. Physiol., A 169, 479-491 (1991).
[CrossRef]

Groot, C.

C. Groot, "On the orientation of young sockeye salmon (Oncorhynchus nerka), during their seaward migration out of lakes," Behaviour 14, 198 (1965).

Guinn, J. J. A.

G. W. Kattawar, G. N. Plass, and J. J. A. Guinn, "Monte Carlo calculations of the polarization of radiation in the Earth's atmosphere-ocean system," J. Phys. Oceanogr. 3, 353-372 (1973).
[CrossRef]

Hagen, R.

N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
[CrossRef] [PubMed]

Hanlon, R. T.

N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
[CrossRef] [PubMed]

N. Shashar, R. T. Hanlon, and A. D. Petz, "Polarization vision helps detect transparent prey," Nature 393, 222-223 (1998).
[CrossRef]

Hawryshyn, C. W.

I. Novales Flamarique and C. W. Hawryshyn, "Is the use of underwater polarized light by fish restricted to crepuscular time periods?" Vision Res. 37, 975-989 (1997).
[CrossRef] [PubMed]

C. W. Hawryshyn, "Polarization vision in fish," Am. Sci. 80, 479-491 (1992).

C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
[CrossRef]

Hobson, E. S.

E. S. Hobson, "Trophic relationships of fishes specialized to feed on zooplankters above coral reefs," in The Ecology of Fishes on Coral Reefs (Academic, 1991), pp. 69-94.

Hojerslev, N. K.

Horch, K. W.

R. B. Forward, K. W. Horch, and T. H. Waterman, "Visual orientation at the water surface by the teleost Zenarchopterus," Biol. Bull. 143, 112-126 (1972).
[CrossRef]

Horvath, G.

G. Horvath and D. Varjú, Polarized Light in Animal Vision: Polarization Patterns in Nature (Springer-Verlag, 2004).

Horváth, G.

S. Sabbah, A. Barta, J. Gal, G. Horváth, and N. Shashar, "Experimental and theoretical study of skylight polarization transmitted through Snell's window of a flat water surface," J. Opt. Soc. Am. A 23, 1978-1988 (2006).
[CrossRef]

G. Horváth and D. Varjú, "Underwater refraction-polarization patterns of skylight perceived by aquatic animals through Snell's window of the flat water surface," Vision Res. 35, 1651-1666 (1995).
[CrossRef] [PubMed]

Ivanoff, A.

A. Ivanoff, "Polarization measurements in the sea," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 151-175.

A. Ivanoff and T. H. Waterman, "Factors, mainly depth and wavelength, affecting the degree of underwater light polarization," J. Mar. Res. 16, 283-307 (1958).

A. Ivanoff and T. H. Waterman, "Elliptical polarization of submarine illumination," J. Mar. Res. 16, 255-282 (1958).

Kattawar, G. W.

J. T. Adams and G. W. Kattawar, "Neutral points in an atmosphere-ocean system. 1: Upwelling light field," Appl. Opt. 36, 1976-1986 (1997).
[CrossRef] [PubMed]

G. W. Kattawar, G. N. Plass, and J. J. A. Guinn, "Monte Carlo calculations of the polarization of radiation in the Earth's atmosphere-ocean system," J. Phys. Oceanogr. 3, 353-372 (1973).
[CrossRef]

Kleerekoper, H.

H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
[CrossRef]

Lerner, A.

S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.

Lundgren, B.

Matis, J. H.

H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
[CrossRef]

McAlary, F. A.

W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.

McFarland, W. N.

W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.

W. N. McFarland, "Light in the sea—correlations with behaviors of fishes and invertebrates," Am. Zool. 26, 389-401 (1986).

Mobley, C. D.

C. D. Mobley, Light and Water (Academic, 1994).

Nicol, J. A. C.

E. J. Denton and J. A. C. Nicol, "Polarization of light reflected from the silvery exterior of the bleak Alburnus alburnus," J. Mar. Biol. Assoc. U.K. 45, 705-709 (1965).
[CrossRef]

Novales Flamarique, I.

I. Novales Flamarique and H. I. Browman, "Foraging and prey-search behaviour of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).

I. Novales Flamarique and C. W. Hawryshyn, "Is the use of underwater polarized light by fish restricted to crepuscular time periods?" Vision Res. 37, 975-989 (1997).
[CrossRef] [PubMed]

Petz, A. D.

N. Shashar, R. T. Hanlon, and A. D. Petz, "Polarization vision helps detect transparent prey," Nature 393, 222-223 (1998).
[CrossRef]

Plass, G. N.

G. W. Kattawar, G. N. Plass, and J. J. A. Guinn, "Monte Carlo calculations of the polarization of radiation in the Earth's atmosphere-ocean system," J. Phys. Oceanogr. 3, 353-372 (1973).
[CrossRef]

Ritz, D. A.

D. A. Ritz, "Polarized-light responses in the shrimp Palaemonetes vulgari (Say)," J. Exp. Mar. Biol. Ecol. 154, 245-250 (1991).
[CrossRef]

Sabbah, S.

S. Sabbah and N. Shashar, "Underwater light polarization and radiance fluctuations induced by surface waves," Appl. Opt. 45, 4726-4739 (2006).
[CrossRef] [PubMed]

S. Sabbah, A. Barta, J. Gal, G. Horváth, and N. Shashar, "Experimental and theoretical study of skylight polarization transmitted through Snell's window of a flat water surface," J. Opt. Soc. Am. A 23, 1978-1988 (2006).
[CrossRef]

S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.

N. Shashar, S. Sabbah, and T. W. Cronin, "Transmission of linearly polarized light in sea water implications for polarization signaling," J. Exp. Biol. 207, 3619-3628 (2004).
[CrossRef] [PubMed]

Schwind, R.

R. Schwind, "Daphnia pulex swims towards the most strongly polarized light—a response that leads to 'shore flight'," J. Exp. Biol. 202, 3631-3635 (1999).
[PubMed]

Shashar, N.

S. Sabbah and N. Shashar, "Underwater light polarization and radiance fluctuations induced by surface waves," Appl. Opt. 45, 4726-4739 (2006).
[CrossRef] [PubMed]

S. Sabbah, A. Barta, J. Gal, G. Horváth, and N. Shashar, "Experimental and theoretical study of skylight polarization transmitted through Snell's window of a flat water surface," J. Opt. Soc. Am. A 23, 1978-1988 (2006).
[CrossRef]

S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.

N. Shashar, S. Sabbah, and T. W. Cronin, "Transmission of linearly polarized light in sea water implications for polarization signaling," J. Exp. Biol. 207, 3619-3628 (2004).
[CrossRef] [PubMed]

T. W. Cronin and N. Shashar, "The linearly polarized light field in clear, tropical marine waters: spatial and temporal variation of light intensity, degree of polarization and e-vector angle," J. Exp. Biol. 204, 2461-2467 (2001).
[PubMed]

N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
[CrossRef] [PubMed]

N. Shashar, R. T. Hanlon, and A. D. Petz, "Polarization vision helps detect transparent prey," Nature 393, 222-223 (1998).
[CrossRef]

Suchanek, T. H.

W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.

Timms, A. M.

H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
[CrossRef]

Timofeyeva, V. A.

V. A. Timofeyeva, "Optics of turbid waters (results of laboratory studies)," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 177-219.

V. A. Timofeyeva, "The degree of polarization of light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 6, 513-522 (1970).

V. A. Timofeyeva, "Plane of vibrations of polarized light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 5, 1049-1057 (1969).

Umminger, B. L.

B. L. Umminger, "Polarotaxis in copepods. I. An endogenous rhythm in polarotaxis in Cyclops vernalis and its relation to vertical migration," Biol. Bull. 135, 239-251 (1968).
[CrossRef]

Varjú, D.

G. Horvath and D. Varjú, Polarized Light in Animal Vision: Polarization Patterns in Nature (Springer-Verlag, 2004).

G. Horváth and D. Varjú, "Underwater refraction-polarization patterns of skylight perceived by aquatic animals through Snell's window of the flat water surface," Vision Res. 35, 1651-1666 (1995).
[CrossRef] [PubMed]

Wahl, C. M.

W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.

Waterman, T. H.

T. H. Waterman, "Polarization sensitivity," in Comparative Physiology and Evolution of Vision in Invertebrates. B: Invertebrate Visual Centers and Behavior I, H.Autrum, ed. (Springer-Verlag, 1981), pp. 281-469.

R. B. Forward and T. H. Waterman, "Evidence for e-vector and light intensity pattern discrimination by the teleost Demogenys," J. Comp. Physiol., A 87, 189-202 (1973).
[CrossRef]

R. B. Forward, K. W. Horch, and T. H. Waterman, "Visual orientation at the water surface by the teleost Zenarchopterus," Biol. Bull. 143, 112-126 (1972).
[CrossRef]

A. Ivanoff and T. H. Waterman, "Factors, mainly depth and wavelength, affecting the degree of underwater light polarization," J. Mar. Res. 16, 283-307 (1958).

A. Ivanoff and T. H. Waterman, "Elliptical polarization of submarine illumination," J. Mar. Res. 16, 255-282 (1958).

T. H. Waterman and W. E. Westell, "Quantitative effect of the sun's position on submarine light polarization," J. Mar. Res. 15, 149-169 (1956).

T. H. Waterman, "Polarization patterns in submarine illumination," Science 120, 927-932 (1954).
[CrossRef] [PubMed]

Wehner, R.

R. Wehner, "Polarization vision—a uniform sensory capacity?" J. Exp. Biol. 204, 2589-2596 (2001).
[PubMed]

Westell, W. E.

T. H. Waterman and W. E. Westell, "Quantitative effect of the sun's position on submarine light polarization," J. Mar. Res. 15, 149-169 (1956).

Wolff, L. B.

L. B. Wolff and A. G. Andreou, "Polarization camera sensors," Image Vis. Comput. 13, 497-509 (1995).
[CrossRef]

Am. Sci.

C. W. Hawryshyn, "Polarization vision in fish," Am. Sci. 80, 479-491 (1992).

Am. Zool.

W. N. McFarland, "Light in the sea—correlations with behaviors of fishes and invertebrates," Am. Zool. 26, 389-401 (1986).

Appl. Opt.

Behaviour

C. Groot, "On the orientation of young sockeye salmon (Oncorhynchus nerka), during their seaward migration out of lakes," Behaviour 14, 198 (1965).

Biol. Bull.

R. B. Forward, K. W. Horch, and T. H. Waterman, "Visual orientation at the water surface by the teleost Zenarchopterus," Biol. Bull. 143, 112-126 (1972).
[CrossRef]

B. L. Umminger, "Polarotaxis in copepods. I. An endogenous rhythm in polarotaxis in Cyclops vernalis and its relation to vertical migration," Biol. Bull. 135, 239-251 (1968).
[CrossRef]

Image Vis. Comput.

L. B. Wolff and A. G. Andreou, "Polarization camera sensors," Image Vis. Comput. 13, 497-509 (1995).
[CrossRef]

Izv., Acad. Sci., USSR, Atmos. Oceanic Phys.

V. A. Timofeyeva, "The degree of polarization of light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 6, 513-522 (1970).

V. A. Timofeyeva, "Plane of vibrations of polarized light in turbid media," Izv., Acad. Sci., USSR, Atmos. Oceanic Phys. 5, 1049-1057 (1969).

J. Comp. Physiol., A

S. M. Goddard and R. B. Forward, "The role of the underwater polarized light pattern, in sun compass navigation of the grass shrimp, Palaemonetes vulgaris," J. Comp. Physiol., A 169, 479-491 (1991).
[CrossRef]

R. B. Forward and T. H. Waterman, "Evidence for e-vector and light intensity pattern discrimination by the teleost Demogenys," J. Comp. Physiol., A 87, 189-202 (1973).
[CrossRef]

H. Kleerekoper, J. H. Matis, A. M. Timms, and P. Gensler, "Locomotor response of the goldfish to polarized light and its e-vector," J. Comp. Physiol., A 86, 27-36 (1973).
[CrossRef]

C. W. Hawryshyn, M. G. Arnold, D. Bowering, and R. L. Cole, "Spatial orientation of rainbow trout to plane-polarized light: the ontogeny of E-vector discrimination and spectral sensitivity characteristics," J. Comp. Physiol., A 166, 565-574 (1990).
[CrossRef]

J. Exp. Biol.

R. Schwind, "Daphnia pulex swims towards the most strongly polarized light—a response that leads to 'shore flight'," J. Exp. Biol. 202, 3631-3635 (1999).
[PubMed]

R. Wehner, "Polarization vision—a uniform sensory capacity?" J. Exp. Biol. 204, 2589-2596 (2001).
[PubMed]

I. Novales Flamarique and H. I. Browman, "Foraging and prey-search behaviour of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).

N. Shashar, S. Sabbah, and T. W. Cronin, "Transmission of linearly polarized light in sea water implications for polarization signaling," J. Exp. Biol. 207, 3619-3628 (2004).
[CrossRef] [PubMed]

T. W. Cronin and N. Shashar, "The linearly polarized light field in clear, tropical marine waters: spatial and temporal variation of light intensity, degree of polarization and e-vector angle," J. Exp. Biol. 204, 2461-2467 (2001).
[PubMed]

J. Exp. Mar. Biol. Ecol.

D. A. Ritz, "Polarized-light responses in the shrimp Palaemonetes vulgari (Say)," J. Exp. Mar. Biol. Ecol. 154, 245-250 (1991).
[CrossRef]

J. Mar. Biol. Assoc. U.K.

E. J. Denton and J. A. C. Nicol, "Polarization of light reflected from the silvery exterior of the bleak Alburnus alburnus," J. Mar. Biol. Assoc. U.K. 45, 705-709 (1965).
[CrossRef]

J. Mar. Res.

A. Ivanoff and T. H. Waterman, "Factors, mainly depth and wavelength, affecting the degree of underwater light polarization," J. Mar. Res. 16, 283-307 (1958).

T. H. Waterman and W. E. Westell, "Quantitative effect of the sun's position on submarine light polarization," J. Mar. Res. 15, 149-169 (1956).

A. Ivanoff and T. H. Waterman, "Elliptical polarization of submarine illumination," J. Mar. Res. 16, 255-282 (1958).

J. Opt. Soc. Am. A

J. Phys. Oceanogr.

G. W. Kattawar, G. N. Plass, and J. J. A. Guinn, "Monte Carlo calculations of the polarization of radiation in the Earth's atmosphere-ocean system," J. Phys. Oceanogr. 3, 353-372 (1973).
[CrossRef]

Nature

N. Shashar, R. T. Hanlon, and A. D. Petz, "Polarization vision helps detect transparent prey," Nature 393, 222-223 (1998).
[CrossRef]

Science

T. H. Waterman, "Polarization patterns in submarine illumination," Science 120, 927-932 (1954).
[CrossRef] [PubMed]

Vision Res.

I. Novales Flamarique and C. W. Hawryshyn, "Is the use of underwater polarized light by fish restricted to crepuscular time periods?" Vision Res. 37, 975-989 (1997).
[CrossRef] [PubMed]

G. Horváth and D. Varjú, "Underwater refraction-polarization patterns of skylight perceived by aquatic animals through Snell's window of the flat water surface," Vision Res. 35, 1651-1666 (1995).
[CrossRef] [PubMed]

N. Shashar, R. Hagen, J. G. Boal, and R. T. Hanlon, "Cuttlefish use polarization sensitivity in predation on silvery fish," Vision Res. 40, 71-75 (2000).
[CrossRef] [PubMed]

Other

W. N. McFarland, C. M. Wahl, T. H. Suchanek, and F. A. McAlary, "The behavior of animals around twilight with emphasis on coral reef communities," in Adaptive Mechanisms in Ecology of Vision, S.N.Archer, ed. (Kluwer Academic, 1999), pp. 583-628.

T. H. Waterman, "Polarization sensitivity," in Comparative Physiology and Evolution of Vision in Invertebrates. B: Invertebrate Visual Centers and Behavior I, H.Autrum, ed. (Springer-Verlag, 1981), pp. 281-469.

S. Sabbah, A. Lerner, C. Erlick, and N. Shashar, "Under water polarization vision—a physical examination," in Recent Research Developments in Experimental & Theoretical Biology (Transworld Research Network, 2005), pp. 123-176.

C. D. Mobley, Light and Water (Academic, 1994).

E. Boss, School of Marine Sciences, University of Maine, Orono, Maine, 04469 (personal communication, 2005).

A. Ivanoff, "Polarization measurements in the sea," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 151-175.

V. A. Timofeyeva, "Optics of turbid waters (results of laboratory studies)," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 177-219.

E. S. Hobson, "Trophic relationships of fishes specialized to feed on zooplankters above coral reefs," in The Ecology of Fishes on Coral Reefs (Academic, 1991), pp. 69-94.

G. Horvath and D. Varjú, Polarized Light in Animal Vision: Polarization Patterns in Nature (Springer-Verlag, 2004).

W. G. Egan, Photometry and Polarization in Remote Sensing (Elsevier, 1985).

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Figures (4)

Fig. 1
Fig. 1

Definitions of light directionality. Solar zenith angle ( θ s ) —the vertical angle between the zenith and the Sun as viewed from outside the water, ranging from 0 ° when the Sun is at the zenith, through 90 ° when the Sun is at the horizon, to angles greater than 90 ° when the Sun is below the horizon. The vertical angle between the refracted light beam and the zenith ( θ r ) ranges from 0 ° when the Sun is at the zenith to 48.6 ° when the Sun is at the horizon, to larger angles when the Sun is below the horizon. The detector zenith angle ( θ p ; also referred to as the viewing zenith angle) is the vertical angle between the zenith and the detector and ranges from 0 ° when the detector is facing the zenith, to 90 ° when the detector is facing horizontally, to 180 ° when the detector is pointing toward the nadir. The solar and detector azimuthal angles ( φ s ) and ( φ p ) are the horizontal angles between the north direction and the Sun or the detector, respectively (the azimuthal angles are measured clockwise from the north when looking downward). The viewing azimuthal angle ( φ ) is the horizontal angle between the two vertical planes containing the Sun and the detector.

Fig. 2
Fig. 2

Qualitative illustration of the e-vector orientation and percent polarization at (a) sunrise and (b) midday. Data are based on measurements and calculations corresponding to the green spectral region ( 500 nm ) . The polarization state is presented for five viewing azimuthal angles ( 0 ° , 45 ° , 90 ° , 135 ° , and 180 ° ) and four viewing zenith angles ( 0 ° , 30 ° , 70 ° , and 90 ° ). The shaded circle represents the theoretical location of Snell’s window. The e-vector orientation is indicated by the tilt of the arrow relative to a tangent to the circle passing through that point, and the percent polarization is indicated by the thickness of the arrows. Consistent with the nomenclature previously used, the figure was reconstructed after Waterman [10], Horváth and Varjú [14], and Cronin and Shashar [11].

Fig. 3
Fig. 3

Underwater polarization at sunrise as a function of the viewing azimuthal angle: (a)–(d) e-vector orientation and (e)–(h) percent polarization at viewing zenith angles of 0 ° , 30 ° , 70 ° , and 90 ° , respectively. Within Snell’s window, the polarization at 350 700 nm is presented, whereas, due to low signal-to-noise ratio of the recorded intensities, outside of Snell’s window the polarization is presented at 380 650 nm . The average (symbols) and standard deviation (error bars) of ten consequent polarization measurements, 1 min apart, starting as the Sun peeked above the mountains (solar zenith angles of 85 ° 87.7 ° ) are depicted.

Fig. 4
Fig. 4

(a)–(e) Predicted and measured e-vector orientations and (f)–(j) percent polarization at five viewing azimuthal angles as a function of the solar zenith angle, at a viewing zenith angle of 90 ° . Polarization values are presented at four wavelengths: 380 (black dots), 450 (gray dots), 550 (gray triangles), and 650 nm (open triangles). For each plot, the Rayleigh scattering prediction was calculated (dashed curves). To calculate the predicted percent polarization, P max was set as the maximal percent polarization recorded throughout the day at the relevant viewing azimuthal angle, measured at 550 nm (the middle of the examined spectrum). Solid vertical lines indicate the solar zenith angle at the actual sunrise. Generally, at large solar zenith angles ( > 75 ° ) , the prediction considerably diverged from the measured polarization.

Equations (3)

Equations on this page are rendered with MathJax. Learn more.

tan ψ = tan θ r sin ( φ s φ p ) sin θ p tan θ r cos θ p cos ( φ s φ p ) ,
P = P max sin 2 Θ sca 1 + cos 2 Θ sca ,
Θ sca = arccos [ cos ( θ r ) cos ( θ p ) + sin ( θ r ) sin ( θ p ) cos ( φ p φ s ) ] .

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