Abstract

The underwater light field is an ever-changing environment. Surface waves induce variability in the radiance and the light's polarization. We examined the dependence of the polarization fluctuations associated with diffuse light (not including contribution from direct skylight) on the viewing zenith angle (30°, 70°, and 90°), solar zenith angle (23°–72°), depth of 0.5–3  m, and light wavelength (380650  nm) while observing within the azimuthal plane in the wind–wave direction. Polarization and radiance fluctuated with time. Light variability (presented by the coefficient of variation calculated over a series of fluctuations in the radiance and percent polarization, and by the standard deviation calculated over a series of fluctuations in the e-vector orientation) was highest at a viewing zenith angle of 70°, depended positively on the solar zenith angle, and decreased with depth at viewing zenith angles of 30° and 70°. Additionally, the variability of the percent polarization was significantly higher than that of the radiance. The temporal light fluctuations offer possibilities, such as enhancing the detection of transparent and reflecting objects; however, they set constraints on the optimal underwater polarization vision by both animals and by the use of instruments.

© 2006 Optical Society of America

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2004 (1)

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]

2001 (5)

H. H. Tynes, G. W. Kattawar, E. P. Zege, I. L. Katsev, A. S. Prikhach, and L. I. Chaikovskaya, "Monte Carlo and multicomponent approximation methods for vector radiative transfer by use of effective Mueller matrix calculations," Appl. Opt. 40, 400-412 (2001).
[CrossRef]

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 behavior of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).

S. L. Degner and C. W. Hawryshyn, "Orientation of rainbow trout (Oncorhynchus mykiss) to multiple patches of linearly polarized light," Can. J. Zool. 79, 407-415 (2001).

2000 (2)

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]

V. V. Maximov, "Environmental factors which may have led to the appearance of colour vision," Philos. Trans. R. Soc. London Ser. B 355, 1239-1242 (2000).
[CrossRef]

1999 (1)

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]

1998 (2)

M. Stramska and T. D. Dickey, "Short-term variability of the underwater light field in the oligotrophic ocean in response to surface waves and clouds," Deep-Sea Res. Suppl. 45, 1393-1410 (1998).
[CrossRef]

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

1997 (1)

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]

1995 (2)

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

1994 (1)

H. I. Browman, I. Novales Flamarique, and C. W. Hawryshyn, "Ultraviolet photoreception contributes to prey search behavior in two species of zooplanktivorous fishes," J. Exp. Biol. 186, 187-198 (1994).

1992 (1)

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

1991 (2)

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]

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

1990 (2)

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 characteristics," J. Comp. Physiol. A 166, 565-574 (1990).
[CrossRef]

M. J. Mcfall-Ngai, "Crypsis in the pelagic environment," Am. Zool. 30, 175-188 (1990).

1989 (2)

J. V. Jelley, "Sea waves: their nature, behaviour, and practical importance," Endeavour 13, 148-156 (1989).
[CrossRef]

G. W. Kattawar and C. N. Adams, "Stokes vector calculations of the submarine light field in an atmosphere-ocean with scattering according to a Rayleigh phase matrix: effect of interface refractive index on radiance and polarization," Limnol. Oceanogr. 34, 1453-1472 (1989).
[CrossRef]

1988 (1)

D. Stramski and J. Dera, "On the mechanism for producing flashing light under a wind-disturbed water surface," Oceanologia 25, 5-21 (1988).

1986 (2)

D. Stramski, "Fluctuations of solar irradiance induced by surface waves in the Baltic," Bull. Pol. Acad. Sci. Earth Sci. 34, 333-344 (1986).

J. Dera and D. Stramski, "Maximum effects of sunlight focusing under a wind-disturbed sea surface," Oceanologia 23, 15-42 (1986).

1983 (1)

W. N. McFarland and E. R. Loew, "Wave-produced changes in underwater light and their relations to vision," Environ. Biol. Fish. 8, 173-184 (1983).
[CrossRef]

1981 (1)

A. J. Underwood, "Techniques of analysis of variance in experimental marine ecology," Oceanoger. Biol. Ann. Rev. 19, 513-605 (1981).

1978 (3)

V. G. Yakubenko and V. P. Nikolayev, "Experimental studies of brightness fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 624-628 (1978).

V. P. Nikolayev and V. G. Yakubenko, "Experimental research into the spatial structure on the fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 301-305 (1978).

J. Dera and J. Olszewski, "Experimental study of short-period irradiance fluctuations under an undulated sea surface," Oceanologia 10, 27-49 (1978).

1977 (1)

V. P. Nikolayev and O. I. Prokopov, "The relationship between statistical characteristics of underwater illumination and certain lighting conditions in the sea," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 13, 734-738 (1977).

1975 (1)

V. P. Nikolayev and M. S. Khulapov, "Use of nonstatistical model to explain the mechanism of underwater illumination fluctuation," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 12, 611-613 (1975).

1974 (2)

O. I. Prokopov, V. P. Nikolayev, A. A. Zhil'tsov, and L. M. Nesterenko, "Some results of a study of light-fluctuations in the sea from 'Chernomor' underwater laboratory," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 11, 737-739 (1974).

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

1973 (2)

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]

1972 (2)

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

V. P. Nikolayev, O. I. Prokopov, G. V. Rosenberg, and V. I. Shevernev, "Statistical characteristics of underwater illumination," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 8, 540-544 (1972).

1970 (2)

R. L. Snyder and J. Dera, "Wave-induced light-field fluctuations in the sea," J. Opt. Soc. Am. 60, 1072-1079 (1970).
[CrossRef]

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

1969 (1)

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

1965 (2)

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

1958 (2)

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).

1957 (1)

1956 (1)

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 (1)

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

Adams, C. N.

G. W. Kattawar and C. N. Adams, "Stokes vector calculations of the submarine light field in an atmosphere-ocean with scattering according to a Rayleigh phase matrix: effect of interface refractive index on radiance and polarization," Limnol. Oceanogr. 34, 1453-1472 (1989).
[CrossRef]

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 characteristics," J. Comp. Physiol. A 166, 565-574 (1990).
[CrossRef]

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 characteristics," J. Comp. Physiol. A 166, 565-574 (1990).
[CrossRef]

Browman, H. I.

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

H. I. Browman, I. Novales Flamarique, and C. W. Hawryshyn, "Ultraviolet photoreception contributes to prey search behavior in two species of zooplanktivorous fishes," J. Exp. Biol. 186, 187-198 (1994).

Chaikovskaya, L. I.

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 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]

Degner, S. L.

S. L. Degner and C. W. Hawryshyn, "Orientation of rainbow trout (Oncorhynchus mykiss) to multiple patches of linearly polarized light," Can. J. Zool. 79, 407-415 (2001).

Denny, M. W.

M. W. Denny, Biology and the Mechanics of the Wave-Swept Environment (Princeton Univ. Press, 1988).

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]

Dera, J.

D. Stramski and J. Dera, "On the mechanism for producing flashing light under a wind-disturbed water surface," Oceanologia 25, 5-21 (1988).

J. Dera and D. Stramski, "Maximum effects of sunlight focusing under a wind-disturbed sea surface," Oceanologia 23, 15-42 (1986).

J. Dera and J. Olszewski, "Experimental study of short-period irradiance fluctuations under an undulated sea surface," Oceanologia 10, 27-49 (1978).

R. L. Snyder and J. Dera, "Wave-induced light-field fluctuations in the sea," J. Opt. Soc. Am. 60, 1072-1079 (1970).
[CrossRef]

J. Dera, S. Sagan, and D. Stramski, "Focusing of sunlight by sea surface waves: new measurement results from the Baltic Sea," in Optics of the Air-Sea Interface, L.Estep, ed., Proc. SPIE 1749,65-72 (1992).

Dickey, T. D.

M. Stramska and T. D. Dickey, "Short-term variability of the underwater light field in the oligotrophic ocean in response to surface waves and clouds," Deep-Sea Res. Suppl. 45, 1393-1410 (1998).
[CrossRef]

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 Zenarchpterus," Biol. Bull. 143, 112-126 (1972).
[CrossRef]

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).

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.

S. L. Degner and C. W. Hawryshyn, "Orientation of rainbow trout (Oncorhynchus mykiss) to multiple patches of linearly polarized light," Can. J. Zool. 79, 407-415 (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]

H. I. Browman, I. Novales Flamarique, and C. W. Hawryshyn, "Ultraviolet photoreception contributes to prey search behavior in two species of zooplanktivorous fishes," J. Exp. Biol. 186, 187-198 (1994).

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 characteristics," J. Comp. Physiol. A 166, 565-574 (1990).
[CrossRef]

Horch, K. W.

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

Horváth, G.

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]

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

Ivanoff, A.

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).

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.

Jelley, J. V.

J. V. Jelley, "Sea waves: their nature, behaviour, and practical importance," Endeavour 13, 148-156 (1989).
[CrossRef]

Jerlov, N. G.

N. G. Jerlov, "Significant relationships between optical properties of the sea," in Optical Aspects of Oceanography, N.G.Jerlov and E.S.Nielsen, eds. (Academic, 1974), pp. 77-94.

Katsev, I. L.

Kattawar, G. W.

H. H. Tynes, G. W. Kattawar, E. P. Zege, I. L. Katsev, A. S. Prikhach, and L. I. Chaikovskaya, "Monte Carlo and multicomponent approximation methods for vector radiative transfer by use of effective Mueller matrix calculations," Appl. Opt. 40, 400-412 (2001).
[CrossRef]

G. W. Kattawar and C. N. Adams, "Stokes vector calculations of the submarine light field in an atmosphere-ocean with scattering according to a Rayleigh phase matrix: effect of interface refractive index on radiance and polarization," Limnol. Oceanogr. 34, 1453-1472 (1989).
[CrossRef]

G. W. Kattawar, "Polarization of light in the ocean," in Ocean Optics, R.W.Spinrad, K.L.Carder, and M.J.Perry, eds. (Oxford U. Press, 1994), pp. 202-225.

Khulapov, M. S.

V. P. Nikolayev and M. S. Khulapov, "Use of nonstatistical model to explain the mechanism of underwater illumination fluctuation," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 12, 611-613 (1975).

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]

Loew, E. R.

W. N. McFarland and E. R. Loew, "Wave-produced changes in underwater light and their relations to vision," Environ. Biol. Fish. 8, 173-184 (1983).
[CrossRef]

E. R. Loew, F. A. McAlary, and W. N. McFarland, "Ultraviolet sensitivity in the larvae of two species of marine atherinid fishes," in Zooplankton Sensory Ecology & Physiology, P.H.Lenz, D.K.Hartline, L.E.Purcell, and D.L.Macmillan, eds. (Gordon & Breach, 1996), pp. 195-209.

Lythgoe, J. N.

J. N. Lythgoe, The Ecology of Vision (Oxford U. Press, 1979).

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]

Maximov, V. V.

V. V. Maximov, "Environmental factors which may have led to the appearance of colour vision," Philos. Trans. R. Soc. London Ser. B 355, 1239-1242 (2000).
[CrossRef]

McAlary, F. A.

E. R. Loew, F. A. McAlary, and W. N. McFarland, "Ultraviolet sensitivity in the larvae of two species of marine atherinid fishes," in Zooplankton Sensory Ecology & Physiology, P.H.Lenz, D.K.Hartline, L.E.Purcell, and D.L.Macmillan, eds. (Gordon & Breach, 1996), pp. 195-209.

Mcfall-Ngai, M. J.

M. J. Mcfall-Ngai, "Crypsis in the pelagic environment," Am. Zool. 30, 175-188 (1990).

McFarland, W. N.

W. N. McFarland and E. R. Loew, "Wave-produced changes in underwater light and their relations to vision," Environ. Biol. Fish. 8, 173-184 (1983).
[CrossRef]

E. R. Loew, F. A. McAlary, and W. N. McFarland, "Ultraviolet sensitivity in the larvae of two species of marine atherinid fishes," in Zooplankton Sensory Ecology & Physiology, P.H.Lenz, D.K.Hartline, L.E.Purcell, and D.L.Macmillan, eds. (Gordon & Breach, 1996), pp. 195-209.

Mobley, C. D.

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

Nesterenko, L. M.

O. I. Prokopov, V. P. Nikolayev, A. A. Zhil'tsov, and L. M. Nesterenko, "Some results of a study of light-fluctuations in the sea from 'Chernomor' underwater laboratory," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 11, 737-739 (1974).

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

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]

Nikolayev, V. P.

V. G. Yakubenko and V. P. Nikolayev, "Experimental studies of brightness fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 624-628 (1978).

V. P. Nikolayev and V. G. Yakubenko, "Experimental research into the spatial structure on the fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 301-305 (1978).

V. P. Nikolayev and O. I. Prokopov, "The relationship between statistical characteristics of underwater illumination and certain lighting conditions in the sea," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 13, 734-738 (1977).

V. P. Nikolayev and M. S. Khulapov, "Use of nonstatistical model to explain the mechanism of underwater illumination fluctuation," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 12, 611-613 (1975).

O. I. Prokopov, V. P. Nikolayev, A. A. Zhil'tsov, and L. M. Nesterenko, "Some results of a study of light-fluctuations in the sea from 'Chernomor' underwater laboratory," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 11, 737-739 (1974).

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

V. P. Nikolayev, O. I. Prokopov, G. V. Rosenberg, and V. I. Shevernev, "Statistical characteristics of underwater illumination," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 8, 540-544 (1972).

Novales Flamarique, I.

I. Novales Flamarique and H. I. Browman, "Foraging and prey-search behavior 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]

H. I. Browman, I. Novales Flamarique, and C. W. Hawryshyn, "Ultraviolet photoreception contributes to prey search behavior in two species of zooplanktivorous fishes," J. Exp. Biol. 186, 187-198 (1994).

Olszewski, J.

J. Dera and J. Olszewski, "Experimental study of short-period irradiance fluctuations under an undulated sea surface," Oceanologia 10, 27-49 (1978).

Petz, A. D.

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

Prikhach, A. S.

Prokopov, O. I.

V. P. Nikolayev and O. I. Prokopov, "The relationship between statistical characteristics of underwater illumination and certain lighting conditions in the sea," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 13, 734-738 (1977).

O. I. Prokopov, V. P. Nikolayev, A. A. Zhil'tsov, and L. M. Nesterenko, "Some results of a study of light-fluctuations in the sea from 'Chernomor' underwater laboratory," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 11, 737-739 (1974).

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

V. P. Nikolayev, O. I. Prokopov, G. V. Rosenberg, and V. I. Shevernev, "Statistical characteristics of underwater illumination," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 8, 540-544 (1972).

Rao, P. V.

P. V. Rao, Statistical Research Methods in the Life Science (Duxbury Press, 1998).

Ritz, D. A.

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

Rosenberg, G. V.

V. P. Nikolayev, O. I. Prokopov, G. V. Rosenberg, and V. I. Shevernev, "Statistical characteristics of underwater illumination," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 8, 540-544 (1972).

Sabbah, S.

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]

Sagan, S.

J. Dera, S. Sagan, and D. Stramski, "Focusing of sunlight by sea surface waves: new measurement results from the Baltic Sea," in Optics of the Air-Sea Interface, L.Estep, ed., Proc. SPIE 1749,65-72 (1992).

Schenck, H.

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.

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]

Shevernev, V. I.

V. P. Nikolayev, O. I. Prokopov, G. V. Rosenberg, and V. I. Shevernev, "Statistical characteristics of underwater illumination," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 8, 540-544 (1972).

Snyder, R. L.

Stramska, M.

M. Stramska and T. D. Dickey, "Short-term variability of the underwater light field in the oligotrophic ocean in response to surface waves and clouds," Deep-Sea Res. Suppl. 45, 1393-1410 (1998).
[CrossRef]

Stramski, D.

D. Stramski and J. Dera, "On the mechanism for producing flashing light under a wind-disturbed water surface," Oceanologia 25, 5-21 (1988).

D. Stramski, "Fluctuations of solar irradiance induced by surface waves in the Baltic," Bull. Pol. Acad. Sci. Earth Sci. 34, 333-344 (1986).

J. Dera and D. Stramski, "Maximum effects of sunlight focusing under a wind-disturbed sea surface," Oceanologia 23, 15-42 (1986).

J. Dera, S. Sagan, and D. Stramski, "Focusing of sunlight by sea surface waves: new measurement results from the Baltic Sea," in Optics of the Air-Sea Interface, L.Estep, ed., Proc. SPIE 1749,65-72 (1992).

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, "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).

Tynes, H. H.

Underwood, A. J.

A. J. Underwood, "Techniques of analysis of variance in experimental marine ecology," Oceanoger. Biol. Ann. Rev. 19, 513-605 (1981).

Varjú, D.

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]

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

Walker, R. E.

R. E. Walker, Marine Light Field Statistics (Wiley, 1994).

Waterman, T. H.

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 Zenarchpterus," 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]

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

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]

Yakubenko, V. G.

V. P. Nikolayev and V. G. Yakubenko, "Experimental research into the spatial structure on the fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 301-305 (1978).

V. G. Yakubenko and V. P. Nikolayev, "Experimental studies of brightness fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 624-628 (1978).

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

Zege, E. P.

Zhil'tsov, A. A.

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

O. I. Prokopov, V. P. Nikolayev, A. A. Zhil'tsov, and L. M. Nesterenko, "Some results of a study of light-fluctuations in the sea from 'Chernomor' underwater laboratory," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 11, 737-739 (1974).

Am. Sci. (1)

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

Am. Zool. (1)

M. J. Mcfall-Ngai, "Crypsis in the pelagic environment," Am. Zool. 30, 175-188 (1990).

Appl. Opt. (1)

Behaviour (1)

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

Biol. Bull. (1)

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

Bull. Pol. Acad. Sci. (1)

D. Stramski, "Fluctuations of solar irradiance induced by surface waves in the Baltic," Bull. Pol. Acad. Sci. Earth Sci. 34, 333-344 (1986).

Can. J. Zool. (1)

S. L. Degner and C. W. Hawryshyn, "Orientation of rainbow trout (Oncorhynchus mykiss) to multiple patches of linearly polarized light," Can. J. Zool. 79, 407-415 (2001).

Deep-Sea Res. Suppl. (1)

M. Stramska and T. D. Dickey, "Short-term variability of the underwater light field in the oligotrophic ocean in response to surface waves and clouds," Deep-Sea Res. Suppl. 45, 1393-1410 (1998).
[CrossRef]

Endeavour (1)

J. V. Jelley, "Sea waves: their nature, behaviour, and practical importance," Endeavour 13, 148-156 (1989).
[CrossRef]

Environ. Biol. Fish. (1)

W. N. McFarland and E. R. Loew, "Wave-produced changes in underwater light and their relations to vision," Environ. Biol. Fish. 8, 173-184 (1983).
[CrossRef]

Image Vis. Comput. (1)

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

Izv. (9)

V. P. Nikolayev, O. I. Prokopov, G. V. Rosenberg, and V. I. Shevernev, "Statistical characteristics of underwater illumination," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 8, 540-544 (1972).

O. I. Prokopov, V. P. Nikolayev, A. A. Zhil'tsov, and L. M. Nesterenko, "Some results of a study of light-fluctuations in the sea from 'Chernomor' underwater laboratory," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 11, 737-739 (1974).

V. P. Nikolayev and O. I. Prokopov, "The relationship between statistical characteristics of underwater illumination and certain lighting conditions in the sea," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 13, 734-738 (1977).

V. P. Nikolayev and M. S. Khulapov, "Use of nonstatistical model to explain the mechanism of underwater illumination fluctuation," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 12, 611-613 (1975).

V. G. Yakubenko, V. P. Nikolayev, O. I. Prokopov, A. A. Zhil'tsov, and L. M. Nesterenko, "On the brightness fluctuations of an underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 10, 1009-1013 (1974).

V. G. Yakubenko and V. P. Nikolayev, "Experimental studies of brightness fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 624-628 (1978).

V. P. Nikolayev and V. G. Yakubenko, "Experimental research into the spatial structure on the fluctuations of the underwater light field," Izv. Acad. Sci. USSR Atmos. Oceanic Phys. 14, 301-305 (1978).

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

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

J. Comp. Physiol. A (4)

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 characteristics," J. Comp. Physiol. A 166, 565-574 (1990).
[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]

J. Exp. Biol. (6)

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 behavior of small juvenile rainbow trout (Oncorhynchus mykiss) under polarized light," J. Exp. Biol. 204, 2415-2422 (2001).

H. I. Browman, I. Novales Flamarique, and C. W. Hawryshyn, "Ultraviolet photoreception contributes to prey search behavior in two species of zooplanktivorous fishes," J. Exp. Biol. 186, 187-198 (1994).

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]

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]

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. (1)

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

J. Mar. Biol. Assoc. U. K. (1)

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. (3)

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. (2)

Limnol. Oceanogr. (1)

G. W. Kattawar and C. N. Adams, "Stokes vector calculations of the submarine light field in an atmosphere-ocean with scattering according to a Rayleigh phase matrix: effect of interface refractive index on radiance and polarization," Limnol. Oceanogr. 34, 1453-1472 (1989).
[CrossRef]

Nature (1)

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

Oceanoger. Biol. Ann. Rev. (1)

A. J. Underwood, "Techniques of analysis of variance in experimental marine ecology," Oceanoger. Biol. Ann. Rev. 19, 513-605 (1981).

Oceanologia (3)

J. Dera and J. Olszewski, "Experimental study of short-period irradiance fluctuations under an undulated sea surface," Oceanologia 10, 27-49 (1978).

J. Dera and D. Stramski, "Maximum effects of sunlight focusing under a wind-disturbed sea surface," Oceanologia 23, 15-42 (1986).

D. Stramski and J. Dera, "On the mechanism for producing flashing light under a wind-disturbed water surface," Oceanologia 25, 5-21 (1988).

Philos. Trans. R. Soc. London Ser. B (1)

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Figures (6)

Fig. 1
Fig. 1

Definition of the radiance's directionality. The 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), at 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), at 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), the vertical angle between the zenith and the detector, ranges from 0° (when the detector is facing the zenith), at 90° (when the detector is facing horizontally), to 180° (when the detector is pointing toward the nadir); the solar and the detector azimuthal angles (φ s ) and (φ p ), the horizontal angles between the direction in which the wind is blowing and the Sun or the detector, respectively [following Mobley (Ref. 3); the azimuthal angles are measured clockwise from the direction of the wind when looking downward]; and the viewing azimuthal angle (φ), the horizontal angle between the two vertical planes containing the Sun and the detector.

Fig. 2
Fig. 2

A 30 s time series of high-rate (3.2 Hz) continuous measurements of the (a) raw radiance, (b) % polarization, (c) e-vector orientation. The horizontal line denotes the time averaged value of the parameter in question. The time series was taken at 600 nm, at 2 m depth with the detector facing a viewing zenith angle of 30°, a solar zenith angle of 27°, a viewing azimuthal angle of 178°.

Fig. 3
Fig. 3

Variability of the radiance, % polarization, and e-vector orientation as a function of the viewing zenith angle (30°, 70°, and 90°) and light wavelength (380–650 nm), taken during 10 measurements days.

Fig. 4
Fig. 4

Average (± standard deviation) of the normalized variability of the light parameters as a function of the viewing zenith angle and the light's wavelength. Repetitions were divided into two groups based on the solar zenith angle. The first group (○, n = 4) incorporated repetitions taken at large solar zenith angles (54°–72°) while the second group (●, n = 6) included repetitions performed at small solar zenith angles (23°–43°). Variability values were normalized by division to the average calculated throughout the spectrum. In (a), (b), and (e) the dependence on the light's wavelength was statistically shown to be bimodal (see Section 2 in text).

Fig. 5
Fig. 5

Effects of water depth on the variability of the radiance, % polarization, and e-vector orientation, as a function of the viewing zenith angle and the light wavelength. Each point represents the average (±standard deviation) of the variability of the light parameters during 3 days of measurement at depths of 0.5 m (●), 1 m (○), 2 m (▼), and 3 m (∇). At viewing zenith angles of 30° and 70°, variability was negatively correlated with depth (30°, ANOVA for radiance and % polarization, and ANOVA preceded by quadratic root transformation for e- vector orientation, F 3,56 = 9.38, 8.83, and 26.23 for radiance, % polarization, and e-vector orientation, respectively. p < 0.001 for all cases; 70°, ANOVA preceded by quadratic root transformation, F 3,56 = 14.34, 36.94, and 49.53 for radiance, % polarization, and e-vector orientation, respectively). At a viewing zenith angle of 90° no significant correlation between depth and the variability of the light parameters was found (nonparametric ANOVA, Kruskal–Wallis, H 3,12 = 6.43, 2.07, and 2.69, p > 0.09, 0.5, and 0.4 for radiance, % polarization, and e-vector orientation after excluding 600–650 nm wavelengths, respectively).

Fig. 6
Fig. 6

Coefficient of variation at 500 nm of the radiance fluctuations as a function of detector depth at viewing zenith angles of (a) 30°, (b) 70°, (c) 90°. The surface waves' averaged frequencies equaled 0.59, 0.56, and 1.12 Hz during days 1, 2, and 3, respectively. During days 1 and 2, when the surface waves' average frequencies were nearly similar, the coefficient of variation varied with depth similarly. However, during day 3, when the surface waves' averaged frequency was different, the highest coefficient of variation was attained at different depths.

Tables (3)

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Table 1 Data of Hydrological and Meteorological Conditions During the Experimental and Control Setups a

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Table 2 Correlation Between the Solar Zenith Angle and the Variability of the Light Parameters a

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Table 3 Correlation Between the Surface Waves' Averaged Frequency and the Dominant Frequency of the Fluctuations in Light Parameters a

Equations (4)

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θ = ( 1 2 ) arctan ( L 0 + L 90 2 L 45 L 90 L 0 ) .
I f   ( θ > 90° ) α = θ 90° ,   otherwise  α = θ + 90° .
L = L 0 + L 90 ,
p = 100 ( L 0 L 90 ) 2 + ( 2 L 45 L 90 L 0 ) 2 L 0 + L 90 .

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