Carbon cycling in the upper waters of the Sargasso Sea: II. Numerical simulation of apparent and inherent optical properties

A mathematical framework to incorporate spectral apparent and inherent optical properties into a one-dimensional ecological simulation (EcoSim) of the Sargasso Sea is developed. The simulation includes equations for spectral algal particulate absorption, in the form of 4 functional groups of phytoplankton, colored degradational matter (CDM), in the form of 2 classes of colored dissolved organic carbon (CDOC), and downwelling diffuse attenuation coefficients (Kd’s) at a 5 nm resolution. Particulate absorption responds to changes in phytoplankton species biomass, pigmentation, and photo-adaptation. CDM absorption responds to concentration changes of labile and relict CDOC. Kd’s respond to changes in spectral total absorption, backscattering, and the average cosine of downwelling photons. The spectral bio-optical outputs provide an additional means of validating an ecological simulation. The vertical and seasonal changes in the diffuse attenuation coefficient at wavelengths of 412, 442, 467, 487, 522, and 567 nm compare well with in situ measurements. There appears to be an underestimation of CDM by the EcoSim that is reflected in the simulated absorption and attenuation at 412 nm. The particulate absorption slope between 412 and 487 nm suggests an overestimation of chlorophyll b and photoprotective carotenoid concentrations from Prochlorococcus functional groups. Simulated Kd(522) and Kd(567) appear to be higher than observations and may result from the exclusion of the inelastic scattering process (i.e. Raman scattering and CDM fluorescence). Results suggest that CDM absorption does not co-vary with particulate absorption. Comparison with a six year time-series of CZCS data suggests that CDM interference of the estimated CZCS chlorophyll a may have been pre-valent in the late spring and the early fall.

[1]  J. Walsh,et al.  CO2 cycling in the coastal ocean. I – A numerical analysis of the southeastern Bering Sea with applications to the Chukchi Sea and the northern Gulf of Mexico , 1994 .

[2]  M. Perry,et al.  Estimating primary production at depth from remote sensing. , 1996, Applied optics.

[3]  R. Goericke,et al.  Chlorophylls a and b and divinyl chlorophylls a and b in the open subtropical North Atlantic Ocean , 1993 .

[4]  R. G. Barlow,et al.  Photosynthetic characteristics of phycoerythrin-containing marine Synechococcus spp. , 1985 .

[5]  R. Goericke,et al.  The marine prochlorophyte Prochlorococcus contributes significantly to phytoplankton biomass and primary production in the Sargasso Sea , 1993 .

[6]  Trevor Platt,et al.  The spectral irradiance field at the surface and in the interior of the ocean: A model for applications in oceanography and remote sensing , 1988 .

[7]  Karen S. Baker,et al.  Oceanic primary production estimates from measurements of spectral irradiance and pigment concentrations , 1987 .

[8]  E. Peltzer,et al.  The use of in situ and airborne fluorescence measurements to determine UV absorption coefficients and DOC concentrations in surface waters , 1995 .

[9]  Josef M. Oberhuber,et al.  An Atlas Based on the COADS Data Set: the Budgets of Heat Buoyancy and Turbulent Kinetic Energy at t , 1988 .

[10]  R. Zepp,et al.  Comparison of photochemical behavior of various humic substances in water: III. Spectroscopic properties of humic substances , 1981 .

[11]  M. Perry,et al.  Quantum yield, relative specific absorption and fluorescence in nitrogen-limited Chaetoceros gracilis , 1987 .

[12]  P. Glibert,et al.  Different patterns of growth and nitrogen uptake in two clones of marineSynechococcus spp. , 1990 .

[13]  Kendall L. Carder,et al.  Carbon cycling in the upper waters of the Sargasso Sea: I. Numerical simulation of differential carbon and nitrogen fluxes , 1999 .

[14]  Nicolas Hoepffner,et al.  Determination of the major groups of phytoplankton pigments from the absorption spectra of total particulate matter , 1993 .

[15]  B. Mitchell,et al.  Microphotometric analysis of individual particle absorption spectra1 , 1988 .

[16]  T. Piatt,et al.  A mechanistic model of photoadaptation in microalgae , 1986 .

[17]  K. Carder,et al.  Marine humic and fulvic acids: Their effects on remote sensing of ocean chlorophyll , 1989 .

[18]  S. Sathyendranath,et al.  Bio-optical characteristics of coastal waters: Absorption spectra of phytoplankton and pigment distribution in the western North Atlantic , 1992 .

[19]  K. Baker,et al.  Evidence for phytoplankton succession and chromatic adaptation in the Sargasso Sea during spring 1985 , 1990 .

[20]  S. Sathyendranath,et al.  Effect of pigment composition on absorption properties of phytoplankton , 1991 .

[21]  H. Gordon,et al.  Remote Assessment of Ocean Color for Interpretation of Satellite Visible Imagery: A Review , 1983 .

[22]  A. Wood,et al.  Biomass of bacteria, cyanobacteria, prochlorophytes and photosynthetic eukaryotes in the Sargasso Sea , 1992 .

[23]  H. Ducklow,et al.  Growth of bacterioplankton and consumption of dissolved organic carbon in the Sargasso Sea , 1996 .

[24]  T. Kana,et al.  Nitrogen starvation in marine Synechococcus strains: clonal differences in phycobiliprotein breakdown and energy coupling , 1992 .

[25]  A. Wood,et al.  Vertical distribution of North Atlantic ultraphytoplankton: analysis by flow cytometry and epifluorescence microscopy , 1988 .

[26]  Dale A. Kiefer,et al.  A two‐component description of spectral absorption by marine particles , 1989 .

[27]  David A. Siegel,et al.  Seasonal variability of light availability and utilization in the Sargasso Sea , 1995 .

[28]  P. B. Wright.  An atlas based on the 'Coads' data set: Fields of mean wind, cloudiness and humidity at the surface of global ocean , 1988 .

[29]  J. Paul,et al.  Relationships between chlorophyll and ocean color constituents as they affect remote‐sensing reflectance models1 , 1986 .

[30]  K. Mopper,et al.  Photochemical source of biological substrates in sea water: implications for carbon cycling , 1989, Nature.

[31]  André Morel,et al.  Relation between total quanta and total energy for aquatic photosynthesis1 , 1974 .

[32]  R. Olson,et al.  Analysis of Synechococcus pigment types in the sea using single and dual beam flow cytometry , 1988 .

[33]  K. Mopper,et al.  Hydroxyl radical photoproduction in the sea and its potential impact on marine processes. , 1990, Science.

[34]  David A. Siegel,et al.  Solar radiation, phytoplankton pigments and the radiant heating of the equatorial Pacific warm pool , 1995 .

[35]  Sol Hellerman,et al.  Normal Monthly Wind Stress Over the World Ocean with Error Estimates , 1983 .

[36]  K. Baker,et al.  Optical properties of the clearest natural waters (200-800 nm). , 1981, Applied optics.

[37]  J. Walsh.  A spatial simulation model of the Peru upwelling ecosystem , 1975 .

[38]  H. Claustre,et al.  Variability in the chlorophyll‐specific absorption coefficients of natural phytoplankton: Analysis and parameterization , 1995 .

[39]  D. Siegel,et al.  Quantification of non-algal light attenuation in the Sargasso Sea: Implications for biogeochemistry and remote sensing , 1996 .

[40]  Francis T. Haxo,et al.  PHOTOSYNTHETIC ACTION SPECTRUM OF THE COCCOLITHOPHORID, EMILIANIA HUXLEYI (HAPTOPHYCEAE): 19′ HEXANOYLOXYFUCOXANTHIN AS ANTENNA PIGMENT 1, 2 , 1985 .

[41]  Frank E. Hoge,et al.  Inherent optical properties of the ocean: Retrieval of the absorption coefficient of chromophoric dissolved organic matter from fluorescence measurements , 1993 .

[42]  K. Hayase,et al.  Sedimentary humic acid and fulvic acid as fluorescent organic materials , 1985 .

[43]  L. Tranvik,et al.  Enhanced bacterial growth in response to photochemical transformation of dissolved organic matter , 1995 .

[44]  K. Carder,et al.  A simple spectral solar irradiance model for cloudless maritime atmospheres , 1990 .

[45]  Motoaki Kishino,et al.  Light Utilization Efficiency and Quantum Yield of Phytoplankton in a Thermally Stratified Seai , 1986 .

[46]  Trevor Platt,et al.  Mathematical formulation of the relationship between photosynthesis and light for phytoplankton , 1976 .

[47]  Dake Chen,et al.  The late summer vertical nutrient mixing in Long Island Sound , 1988 .

[48]  D. Vaulot,et al.  Photosynthetic picoplankton community structure in the subtropical North Pacific Ocean near Hawaii (station ALOHA) , 1993 .

[49]  D. Siegel,et al.  Concurrent high resolution bio‐optical and physical time series observations in the Sargasso Sea during the spring of 1987 , 1991 .

[50]  E. Druffel,et al.  Radiocarbon in dissolved organic matter in the central North Pacific Ocean , 1987, Nature.

[51]  K. S. Baker,et al.  Optical variability and pigment biomass in the Sargasso Sea as determined using deep-sea optical mooring data , 1991 .

[52]  A. Morel.  Optical modeling of the upper ocean in relation to its biogenous matter content (case I waters) , 1988 .

[53]  R. Zepp,et al.  Photochemical production of dissolved inorganic carbon from terrestrial organic matter: significance to the oceanic organic carbon cycle , 1995 .

[54]  Robert R. Bidigare,et al.  Seasonal patterns of ocean biogeochemistry at the U.S. JGOFS Bermuda Atlantic time-series study site , 1994 .

[55]  André Morel,et al.  Light and marine photosynthesis: a spectral model with geochemical and climatological implications , 1991 .

[56]  R Iturriaga,et al.  Temporal and spatial variability of chroococcoid cyanobacteria Synechococcus spp. specific growth rates and their contribution to primary production in the Sargasso Sea , 1988 .

[57]  Dale A. Kiefer,et al.  In-vivo absorption properties of algal pigments , 1990, Defense, Security, and Sensing.

[58]  Mary Ann Moran,et al.  Photochemical release of biologically available nitrogen from aquatic dissolved organic matter , 1996, Nature.

[59]  S. W. Jeffrey.  THE OCCURRENCE OF CHLOROPHYLL C1 AND C2 IN ALGAE 1 , 1976 .

[60]  Ronald D. Jones,et al.  Photochemical degradation of dissolved organic carbon and its impact on the oceanic carbon cycle , 1991, Nature.

[61]  P. Falkowski,et al.  A simulation model of the effects of vertical mixing on primary productivity , 1981 .

[62]  K. Carder,et al.  Reflectance Model for Quantifying Chlorophyll- a in the Presence of Productivity Degradation Products , 1991 .

[63]  Daniel Vaulot,et al.  The importance of Prochlorococcus to community structure in the central North Pacific Ocean , 1994 .

[64]  M. D. Keller,et al.  COCCOID EUKARYOTIC MARINE ULTRAPLANKTERS WITH FOUR DIFFERENT HPLC PIGMENT SIGNATURES 1 , 1988 .

[65]  K. Carder,et al.  A remote‐sensing reflectance model of a red‐tide dinoflagellate off west Florida1 , 1985 .

[66]  Sarah A. Green,et al.  Optical absorption and fluorescence properties of chromophoric dissolved organic matter in natural waters , 1994 .

[67]  S. Gorshkov,et al.  World ocean atlas , 1976 .

[68]  William K. W. Li.  Primary production of prochlorophytes, cyanobacteria, and eucaryotic ultraphytoplankton: Measurements from flow cytometric sorting , 1994 .

[69]  Sallie W. Chisholm,et al.  Comparative physiology of Synechococcus and Prochlorococcus: influence of light and temperature on growth, pigments, fluorescence and absorptive properties , 1995 .

[70]  Craig E. Williamson,et al.  The attenuation of solar UV radiation in lakes and the role of dissolved organic carbon , 1995 .

[71]  R. Bidigare,et al.  Seasonal variability of bio-optical and physical properties in the Sargasso sea , 1993 .

[72]  R. Olson,et al.  Pigments, size, and distributions of Synechococcus in the North Atlantic and Pacific Oceans , 1990 .

[73]  G. Mellor,et al.  Development of a turbulence closure model for geophysical fluid problems , 1982 .

[74]  D. Kiefer,et al.  A steady state description ofgrowth and light absorption in the marine planktonic diatom Skeletonema costatum , 1989 .

[75]  M. D. Keller,et al.  The effects of light quality and intensity on photosynthesis and growth of marine eukaryotic and prokaryotic phytoplankton clones , 1987 .

[76]  K. Carder,et al.  Pigment packaging and Chl a‐specific absorption in high‐light oceanic waters , 1997 .

[77]  P. Santschi,et al.  Dynamics of dissolved organic carbon (DOC) in oceanic environments , 1995 .

[78]  Larry A. Chesal,et al.  The structure of marine fulvic and humic acids , 1983 .

[79]  P. Hatcher,et al.  Natural photolysis by ultraviolet irradiance of recalcitrant dissolved organic matter to simple substrates for rapidbacterial metabolism , 1995 .

[80]  J. Raven,et al.  GROWTH, PHOTOSYNTHESIS AND MAINTENANCE METABOLIC COST IN THE DIATOM PHAEODACTYLUM TRICORNUTUM AT VERY LOW LIGHT LEVELS 1 , 1986 .

[81]  B. Osborne,et al.  Light and Photosynthesis in Aquatic Ecosystems. , 1985 .

[82]  F. Muller‐Karger,et al.  The effects of temporal variability of mixed layer depth on primary productivity around Bermuda , 1994 .

[83]  E. Laws,et al.  Nutrient‐ and light‐limited growth of Thalassiosira fluviatilis in continuous culture, with implications for phytoplankton growth in the ocean , 1980 .

[84]  R. L. Hulstrom,et al.  Terrestrial solar spectral data sets , 1982 .

[85]  James W. Brown,et al.  A semianalytic radiance model of ocean color , 1988 .

[86]  B. Osborne,et al.  Light absorption by a marine diatom: experimental observations and theoretical calculations of the package effect in a small Thalassiosira species , 1987 .

[87]  T. Platt,et al.  Vertical Nitrate Fluxes in the Oligotrophic Ocean , 1986, Science.

[88]  Robin Merl Pope.  Optical Absorption of Pure Water and Sea Water Using the Integrating Cavity Absorption Meter. , 1993 .

[89]  Thomas R. Anderson,et al.  A spectrally averaged model of light penetration and photosynthesis , 1993 .

[90]  R. Olson,et al.  Spatial and temporal distributions of prochlorophyte picoplankton in the North Atlantic Ocean , 1990 .

[91]  K. Hayase,et al.  Sedimentary humic acid and fulvic acid as surface active substances , 1983 .

[92]  S. W. Jeffrey,et al.  EFFECT OF BLUE‐GREEN LIGHT ON PHOTOSYNTHETIC PIGMENTS AND CHLOROPLAST STRUCTURE IN THE MARINE DIATOM STEPHANOPYXIS TURRIS 1 , 1977 .

[93]  M. Perry,et al.  Photoadaption in marine phytoplankton: Response of the photosynthetic unit , 1981 .

[94]  O. Schofield,et al.  Spectral photosynthesis, quantum yield and blue-green light enhancement of productivity rates in the diatom Chaetoceros gracile and the prymnesiophyte Emiliania huxleyi , 1990 .

[95]  D. Vaulot,et al.  Photoacclimation of Prochlorococcus sp. (Prochlorophyta) Strains Isolated from the North Atlantic and the Mediterranean Sea , 1993, Plant physiology.

[96]  M. Perry,et al.  Modeling in situ phytoplankton absorption from total absorption spectra in productive inland marine waters , 1989 .

[97]  P. Glibert,et al.  Effect of irradiances up to 2000 μE m−2 s−1 on marine Synechococcus WH7803—II. Photosynthetic responses and mechanisms , 1987 .

[98]  F. Muller‐Karger,et al.  Meridional fluxes of dissolved organic matter in the north Atlantic Ocean , 1992 .

[99]  H. Ducklow,et al.  Annual flux of dissolved organic carbon from the euphotic zone in the northwestern Sargasso Sea , 1994, Nature.

[100]  K. Mopper,et al.  Formation of carbonyl compounds from UV‐induced photodegradation of humic substances in natural waters: Fate of riverine carbon in the sea , 1990 .

[101]  F. Muller‐Karger,et al.  Analysis of Ocean Color Components Within Stratified and Well-Mixed Waters of the Western English Channel , 1995 .

[102]  Dennis A. Hansell,et al.  Predominance of vertical loss of carbon from surface waters of the equatorial Pacific Ocean , 1997, Nature.