Abstract

The backscattering properties of marine phytoplankton, which are assumed to vary widely with differences in size, shape, morphology and internal structure, have been directly measured in the laboratory on a very limited basis. This work presents results from laboratory analysis of the backscattering properties of thirteen phytoplankton species from five major taxa. Optical measurements include portions of the volume scattering function (VSF) and the absorption and attenuation coefficients at nine wavelengths. The VSF was used to obtain the backscattering coefficient for each species, and we focus on intra- and interspecific variability in spectral backscattering in this work. Ancillary measurements included chlorophyll-a concentration, cell concentration, and cell size, shape and morphology via microscopy for each culture. We found that the spectral backscattering properties of phytoplankton deviate from theory at wavelengths where pigment absorption is significant. We were unable to detect an effect of cell size on the spectral shape of backscattering, but we did find a relationship between cell size and both the backscattering ratio and backscattering cross-section. While particulate backscattering at 555 nm was well correlated to chlorophyll-a concentration for any given species, the relationship was highly variable between species. Results from this work indicate that phytoplankton cells may backscatter light at significantly higher efficiencies than what is predicted by Mie theory, which has important implications for closing the underwater and remotely sensed light budget.

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  58. G. A. Jackson, R. Maffione, D. K. Costello, A. L. Alldredge, B. E. Logan, and H. G. Dam, “Particle size spectra between 1 μm and 1 cm at Monterey Bay determined using multiple instruments,” Deep-Sea Res. 44(11), 1739–1767 (1997), doi:.
    [CrossRef]

2009 (2)

J. M. Sullivan and M. S. Twardowski, “Angular shape of the oceanic particulate volume scattering function in the backward direction,” Appl. Opt. 48(35), 6811–6819 (2009).
[CrossRef] [PubMed]

G. Dall'Olmo, T. K. Westberry, M. J. Behrenfeld, E. Boss, and W. H. Slade, “Significant contribution of large particles to optical backscattering in the open ocean,” Biogeosci. 6(6), 947–967 (2009).
[CrossRef]

2008 (1)

Y. Huot, A. Morel, M. S. Twardowski, D. Stramski, and R. A. Reynolds, “Particle optical backscattering along a chlorophyll gradient in the upper layer of the eastern South Pacific Ocean,” Biogeosci. 5(2), 495–507 (2008).
[CrossRef]

2007 (7)

J.-F. Berthon, E. Shybanov, M. E.-G. Lee, and G. Zibordi, “Measurements and modeling of the volume scattering function in the coastal northern Adriatic Sea,” Appl. Opt. 46(22), 5189–5203 (2007).
[CrossRef] [PubMed]

D. Stramski, M. Babin, and S. B. Woźniak, “Variations in the optical properties of terrigenous mineral-rich particulate matter suspended in seawater,” Limnol. Oceanogr. 52, 2418–2433 (2007).
[CrossRef]

E. Boss, R. Collier, G. Larson, K. Fennel, and W. S. Pegau, “Measurements of spectral optical properties and their relation to biogeochemical variables and processes in Crater Lake, Crater Lake National Park, OR,” Hydrobiologia 574(1), 149–159 (2007).
[CrossRef]

A. L. Whitmire, E. Boss, T. J. Cowles, and W. S. Pegau, “Spectral variability of the particulate backscattering ratio,” Opt. Express 15(11), 7019–7031 (2007).
[CrossRef] [PubMed]

A. Samoc, A. Miniewicz, M. Samoc, and J. G. Grote, “Refractive-index anisotropy and optical dispersion in films of deoxyribonucleic acid,” J. Appl. Polym. Sci. 105(1), 236–245 (2007).
[CrossRef]

M. S. Twardowski, H. Claustre, S. A. Freeman, D. Stramski, and Y. Huot, “Optical backscattering properties of the “clearest” natural waters,” Biogeosci. 4(6), 1041–1058 (2007), www.biogeosciences.net/4/1041/2007 .
[CrossRef]

W. R. Clavano, E. Boss, and L. Karp-Boss, “Inherent Optical Properties of Non-Spherical Marine-Like Particles - From Theory to Observations,” Oceanography and Marine Biology: An Annual Review 45, 1–38 (2007).
[CrossRef]

2006 (2)

T. Dickey, M. Lewis, and G. Chang, “Optical oceanography: Recent advances and future directions using global remote sensing and in situ observations,” Rev. Geophys. 44(1), RG1001 (2006), doi:.
[CrossRef]

J. J. Walsh, J. K. Jolliff, B. P. Darrow, J. M. Lenes, S. P. Milroy, A. Remsen, D. A. Dieterle, K. L. Carder, F. R. Chen, G. A. Vargo, R. H. Weisberg, K. A. Fanning, F. E. Muller-Karger, E. Shinn, K. A. Steidinger, C. A. Heil, C. R. Tomas, J. S. Prospero, T. N. Lee, G. J. Kirkpatrick, T. E. Whitledge, D. A. Stockwell, T. A. Villareal, A. E. Jochens, and P. S. Bontempi, “Red tides in the Gulf of Mexico: Where, when, and why?” J. Geophys. Res. 111(C11C11003), 1–46 (2006).
[CrossRef] [PubMed]

2005 (1)

2004 (4)

E. Boss, W. S. Pegau, M. Lee, M. S. Twardowski, E. Shybanov, G. Korotaev, and F. Baratange, “The particulate backscattering ratio at LEO-15 and its use to study particle composition and distribution,” J. Geophys. Res. 109(C1), C01014 (2004), doi:.
[CrossRef]

R. D. Vaillancourt, C. W. Brown, R. R. L. Guillard, and W. M. Balch, “Light backscattering properties of marine phytoplankton: relationships to cell size, chemical composition, and taxonomy,” J. Plankton Res. 26(2), 191–212 (2004).
[CrossRef]

D. Stramski, E. Boss, D. Bogucki, and K. J. Voss, “The role of seawater constituents in light backscattering in the ocean,” Prog. Oceanogr. 61(1), 27–56 (2004).
[CrossRef]

A. Quirantes and S. Bernard, “Light scattering by marine algae: two-layer spherical and nonspherical models,” J. Quant. Spectrosc. Radiat. Transf. 89(1-4), 311–321 (2004).
[CrossRef]

2003 (2)

M. E. Lee and M. R. Lewis, “A new method for the measurement of the optical volume scattering function in the upper ocean,” J. Atmos. Ocean. Technol. 20(4), 563–671 (2003).
[CrossRef]

D. Stramski and J. Piskozub, “Estimation of scattering error in spectrophotometric measurements of light absorption by aquatic particles from three-dimensional radiative transfer simulations,” Appl. Opt. 42(18), 3634–3646 (2003).
[CrossRef] [PubMed]

2001 (2)

2000 (2)

Y. Bhaud, D. Guillebault, J. Lennon, H. Defacque, M. O. Soyer-Gobillard, and H. Moreau, “Morphology and behaviour of dinoflagellate chromosomes during the cell cycle and mitosis,” J. Cell Sci. 113(Pt 7), 1231–1239 (2000).
[PubMed]

H. Loisel and D. Stramski, “Estimation of the inherent optical properties of natural waters from the irradiance attenuation coefficient and reflectance in the presence of Raman scattering,” Appl. Opt. 39(18), 3001–3011 (2000).
[CrossRef]

1998 (2)

K. Witkowski, T. Krol, A. Zielinski, and E. Kuten, “A light-scattering matrix for unicellular marine phytoplankton,” Limnol. Oceanogr. 43(5), 859–869 (1998).
[CrossRef]

H. Volten, J. F. Haan, J. W. Hovenier, R. Schreurs, W. Vassen, A. G. Dekker, H. J. Hoogenboom, F. Charlton, and R. Wouts, “Laboratory Measurements of Angular Distributions of Light Scattered by Phytoplankton and Silt,” Limnol. Oceanogr. 43(6), 1180–1197 (1998).
[CrossRef]

1997 (5)

W. S. Pegau, D. Gray, and J. R. V. Zaneveld, “Absorption and attenuation of visible and near-infrared light in water: dependence on temperature and salinity,” Appl. Opt. 36(24), 6035–6046 (1997).
[CrossRef] [PubMed]

R. A. Maffione and D. R. Dana, “Instruments and methods for measuring the backward-scattering coefficient of ocean waters,” Appl. Opt. 36(24), 6057–6067 (1997).
[CrossRef] [PubMed]

D. Stramski and C. D. Mobley, “Effects of microbial particles on oceanic optics: A database of single-particle optical properties,” Limnol. Oceanogr. 42(3), 538–549 (1997).
[CrossRef]

G. A. Jackson, R. Maffione, D. K. Costello, A. L. Alldredge, B. E. Logan, and H. G. Dam, “Particle size spectra between 1 μm and 1 cm at Monterey Bay determined using multiple instruments,” Deep-Sea Res. 44(11), 1739–1767 (1997), doi:.
[CrossRef]

R. F. Davis, C. C. Moore, J. R. V. Zaneveld, and J. M. Napp, “Reducing the effects of fouling on chlorophyll estimates derived from long-term deployments of optical instruments,” J. Geophys. Res. 102(C3), 5851–5855 (1997).
[CrossRef]

1996 (1)

E. Aas, “Refractive index of phytoplankton derived from its metabolite composition,” J. Plankton Res. 18(12), 2223–2249 (1996).
[CrossRef]

1994 (1)

J. R. V. Zaneveld, J. C. Kitchen, and C. C. Moore, ““Scattering error correction of reflecting tube absorption meter,” Ocean Optics XII,” Proc. SPIE 2258, 44–55 (1994).
[CrossRef]

1992 (2)

Y.-H. Ahn, A. Bricaud, and A. Morel, “Light backscattering efficiency and related properties of some phytoplankters,” Deep-Sea Res. 39(11–12), 1835–1855 (1992).
[CrossRef]

J. C. Kitchen and J. R. V. Zaneveld, “A three-layered sphere model of the optical properties of phytoplankton,” Limnol. Oceanogr. 37(8), 1680–1690 (1992).
[CrossRef]

1991 (2)

A. Morel and Y.-H. Ahn, “Optics of heterotrophic nanoflagellates and ciliates: A tentative assessment of their scattering role in oceanic waters compared to those of bacterial and algal cells,” J. Mar. Res. 49(1), 177–202 (1991).
[CrossRef]

D. Stramski and D. A. Kiefer, “Light scattering by microorganisms in the open ocean,” Prog. Oceanogr. 28(4), 343–383 (1991).
[CrossRef]

1990 (2)

A. Morel and Y.-H. Ahn, “Optical efficiency factors of free-living marine bacteria: Influence of bacterioplankton upon the optical properties and particulate organic carbon in oceanic waters,” J. Mar. Res. 48(1), 145–175 (1990).
[CrossRef]

T. Oishi, “Significant relationship between the backward scattering coefficient of sea water and the scatterance at 120°,” Appl. Opt. 29(31), 4658–4665 (1990).
[CrossRef] [PubMed]

1989 (1)

M. S. Quinby-Hunt, A. J. Hunt, K. Lofftus, and D. Shapiro, “Polarized-light scattering studies of marine Chlorella,” Limnol. Oceanogr. 34(8), 1587–1600 (1989).
[CrossRef]

1988 (1)

A. Bricaud, A. L. Bedhomme, and A. Morel, “Optical properties of diverse phytoplanktonic species: Experimental results and theoretical interpretation,” J. Plankton Res. 10(5), 851–873 (1988).
[CrossRef]

1986 (2)

A. Bricaud and A. Morel, “Light attenuation and scattering by phytoplanktonic cells: a theoretical modeling,” Appl. Opt. 25(4), 571–580 (1986).
[CrossRef] [PubMed]

A. Morel and A. Bricaud, “Inherent optical properties of algal cells, including picoplankton. Theoretical and experimental results,” Can. Bull. Fish. Aquat. Sci. 214, 521–559 (1986).

1984 (1)

U. J. Santore, “Some aspects of taxonomy in the Cryptophyceae,” New Phytol. 98(4), 627–646 (1984).
[CrossRef]

1983 (1)

A. Bricaud, A. Morel, and L. Prieur, “Optical efficiency factors of some phytoplankters,” Limnol. Oceanogr. 28(5), 816–832 (1983).
[CrossRef]

1981 (1)

A. Morel and A. Bricaud, “Theoretical results concerning light absorption in a discrete medium, and application to specific absorption of phytoplankton,” Deep-Sea Res. 28(11), 1375–1393 (1981).
[CrossRef]

1979 (1)

1975 (1)

1974 (1)

M. A. Faust, “Structure of the periplast of Cryptomonas ovata var. palustris,” J. Phycol. 10, 121–124 (1974).

1963 (1)

C. S. Yentsch and D. W. Menzel, “A method for the determination of phytoplankton chlorophyll and phaeophytin by fluorescence,” Deep-Sea Res. 10, 221–231 (1963).

Aas, E.

E. Aas, “Refractive index of phytoplankton derived from its metabolite composition,” J. Plankton Res. 18(12), 2223–2249 (1996).
[CrossRef]

Ahn, Y.-H.

Y.-H. Ahn, A. Bricaud, and A. Morel, “Light backscattering efficiency and related properties of some phytoplankters,” Deep-Sea Res. 39(11–12), 1835–1855 (1992).
[CrossRef]

A. Morel and Y.-H. Ahn, “Optics of heterotrophic nanoflagellates and ciliates: A tentative assessment of their scattering role in oceanic waters compared to those of bacterial and algal cells,” J. Mar. Res. 49(1), 177–202 (1991).
[CrossRef]

A. Morel and Y.-H. Ahn, “Optical efficiency factors of free-living marine bacteria: Influence of bacterioplankton upon the optical properties and particulate organic carbon in oceanic waters,” J. Mar. Res. 48(1), 145–175 (1990).
[CrossRef]

Alldredge, A. L.

G. A. Jackson, R. Maffione, D. K. Costello, A. L. Alldredge, B. E. Logan, and H. G. Dam, “Particle size spectra between 1 μm and 1 cm at Monterey Bay determined using multiple instruments,” Deep-Sea Res. 44(11), 1739–1767 (1997), doi:.
[CrossRef]

Babin, M.

D. Stramski, M. Babin, and S. B. Woźniak, “Variations in the optical properties of terrigenous mineral-rich particulate matter suspended in seawater,” Limnol. Oceanogr. 52, 2418–2433 (2007).
[CrossRef]

Balch, W. M.

R. D. Vaillancourt, C. W. Brown, R. R. L. Guillard, and W. M. Balch, “Light backscattering properties of marine phytoplankton: relationships to cell size, chemical composition, and taxonomy,” J. Plankton Res. 26(2), 191–212 (2004).
[CrossRef]

Baratange, F.

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Figures (7)

Fig. 1
Fig. 1

Angular variation in χ p for 13 phytoplankton cultures. The solid vertical line is the centroid measuremenet angle of the Hydroscat-6 (140°). A yellow ‘X’ is shown at χ p = 1.21, the mean value over all wavelengths and all cultures sampled. Where three or more wavelengths were used, dashed lines are shown at two standard deviations from the mean χ p value for the culture. The measurement wavelengths (in nm) are shown in the legend above. The cultures shown are: (A) Amphidinium carterae, (B) Ditylum brightwellii, (C) Dunaliella tertiolecta, (D) Emeliana huxleyi, (E) Gephyrocapsa oceanica, (F) Isochrysis galbana, (G) Prorocentrum minimum, (H) Pyramimonas parkeae, (I) Stephanopyxis turris, (J) Thoracosphaera heimii, (K) Thalassiosira pseudonana, (L) Thalassiosira weissflogii, (M) Pycnococcus provasolii, and (N) mean of all cultures at all wavelengths with 95% confidence intervals.

Fig. 2
Fig. 2

Angular variation in mean χp for oceanic particles and phytoplankton cultures (see legend). χp at 555 nm is shown for this study (bold line) with 95% confidence intervals (dotted lines). The other wavelengths used were 532 nm for Boss and Pegau [BP; 26], 550 nm for Chami et al. [Ch; 31], 658 nm for Sullivan and Twardowski. [ST; 34], and 514 nm for Vaillancourt et al. [V; 21].

Fig. 3
Fig. 3

Spectral backscattering coefficients for fifteen phytoplankton species. Error bars represent one standard deviation of the mean of data collected over at least two minutes. (A) Diatom species sampled. Where more than one strain was sampled, the strain number is shown. (B) Flagellates. (C) Dinoflagellates. Data shown at 676 nm are contaminated by fluorescence (denoted with an asterisk). These data are shown for reference only.

Fig. 4
Fig. 4

Spectral particulate backscattering ratios for fifteen phytoplankton cultures, separated into taxonomic groups. Error bars show 15% of the signal at each wavelength. (A) Diatoms. Where more than one species was sampled, the strain number is shown (B) Flagellates (C) Dinoflagellates. Data shown at 676 nm are contaminated by fluorescence (denoted with an asterisk). These data are shown for reference only.

Fig. 5
Fig. 5

The backscattering coefficient at 555 nm x1000 versus chlorophyll-a concentration for culture dilution series experiments. (A) data for all cultures (B) data for selected cultures with chlorophyll concentrations of less than 1.5 mg m−3 (C) data for selected cultures with chlorophyll-a concentrations up to 25 mg m−3.

Fig. 6
Fig. 6

(A) The particulate backscattering ratio at 555 nm versus the equivalent spherical diameter for phytoplankton cultures measured with a Coulter counter or light microscopy. (B) The backscattering cross-section at 555 nm versus the equivalent spherical diameter for phytoplankton cultures. The equations for a power law fit and the R2 of the fits are shown.

Fig. 7
Fig. 7

Spectral values of IOP cross-sections and backscattering ratios for cultures that were in the same size range as those shown by Stramski et. al. (2001). Red lines are our data, black lines are from Stramski et al. (2001). The absorption (A) and scattering cross-sections (B) from Stramski et. al. (2001) were based on measurements. The backscattering cross-section (C) and backscattering ratio (D) were model results in Stramski et al. (2001), and were measured in our study.

Tables (2)

Tables Icon

Table 1 Summary of phytoplankton characteristics. Optical data are shown at 442 nm

Tables Icon

Table 2 Average value and relative standard deviation, σ, of χ p for thirteen phytoplankton species from 90 to 170 degrees at six wavelengths. N is the number of cultures available to calculate the mean and standard deviation

Equations (7)

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b b = 2 π π 2 π β ( θ ) sin θ d θ
b b p ( λ ) = 2 π χ p ( θ ) β p ( λ , θ )
χ p ( λ , θ ) = b b p ( λ ) 2 π β p ( λ , θ )
σ b b = b b p ( λ ) N / V
Q b b = σ b b π × r 2
b bp / b p ( 555 )   =  4 . 39 0 x1 0 3 x ESD 0. 432 ;  R 2 =   0. 56
σ bb ( 555 )   =  4 . 269x1 0 14 x ESD 2 .0 28 ;  R 2 =   0. 84

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