Subsurface chlorophyll maximum layers: Enduring enigma or mystery solved?

Annual Review of Marine Science

Volumn 7, Issue , 2015, Pages 207-239

  Cullen, John J ^a  

^a DALHOUSIE UNIVERSITY   (Canada)

Author keywords

acclimation; behavior; carbon to chlorophyll ratio; Deep chlorophyll maximum; growth; phytoplankton; primary production; stratification; thin layers; turbulence

Indexed keywords

CHLOROPHYLL; ECOLOGY; FORECASTING; GROWTH (MATERIALS); NUTRIENTS; THERMAL STRATIFICATION; TURBULENCE;

ACCLIMATION; BEHAVIOR; CARBON-TO-CHLOROPHYLL RATIOS; DEEP CHLOROPHYLL MAXIMUMS; PRIMARY PRODUCTION; THIN LAYERS;

PHYTOPLANKTON;

BUOYANCY; CHLOROPHYLL; CONCENTRATION (COMPOSITION); ENVIRONMENTAL RESPONSE; PERSISTENCE; STRATIFORM DEPOSIT; VERTICAL MIXING;

BACILLARIOPHYTA; CYANOBACTERIA;

CHLOROPHYLL; SEA WATER;

ACCLIMATIZATION; BIOMASS; CHEMISTRY; ENVIRONMENTAL MONITORING; GROWTH, DEVELOPMENT AND AGING; LIGHT; METABOLISM; MICROBIOLOGY; PHYSIOLOGY; PHYTOPLANKTON; POPULATION DENSITY; PROCEDURES; SEA; STATISTICS AND NUMERICAL DATA; TEMPERATURE; THEORETICAL MODEL;

ACCLIMATIZATION; BIOMASS; CHLOROPHYLL; ENVIRONMENTAL MONITORING; LIGHT; MODELS, THEORETICAL; OCEANS AND SEAS; PHYTOPLANKTON; POPULATION DENSITY; SEAWATER; TEMPERATURE;

EID: 84948614453     PISSN: 19411405     EISSN: 19410611     Source Type: Journal    
DOI: 10.1146/annurev-marine-010213-135111     Document Type: Article

References (156)

1. Anderson GC. 1969. Subsurface chlorophyll maximum in the northeast Pacific Ocean. Limnol. Oceanogr. 14:386-91
2. Babin M, Morel A, Claustre H, Bricaud A, Kolber Z, Falkowski PG. 1996. Nitrogen-and irradiance-dependent variations of the maximum quantum yield of carbon fixation in eutrophic, mesotrophic and oligotrophic marine systems. Deep-Sea Res. I 43:1241-72
3. Balch WM, Bowler BC, Byrne CF. 1997. Sea surface temperature gradients, baroclinicity, and vegetation gradients in the sea. J. Plankton Res. 19:1829-58
4. Banse K. 1987. Clouds, deep chlorophyll maxima and the nutrient supply to the mixed layer of stratified water bodies. J. Plankton Res. 9:1031-36
5. Banse K. 1995. Zooplankton: pivotal role in the control of ocean production. ICES J. Mar. Sci. 52:265-77
6. Banse K. 2004. Should we continue to use the 1% light depth convention for estimating the compensation depth of phytoplankton for another 70 years? Limnol. Oceanogr. Bull. 13:49-52
7. Banse K. 2013. Reflections about chance in my career, and on the top-down regulated world. Annu. Rev. Mar. Sci. 5:1-19
8. Barber RT, Hilting AK. 2002. History of the study of plankton productivity. In Phytoplankton Productivity: Carbon Assimilation in Marine and Freshwater Ecosystems, ed. PJLB Williams, DN Thomas, CS Reynolds, pp. 16-43. Oxford, UK: Blackwell Sci.
9. Beckmann A, Hense I. 2007. Beneath the surface: characteristics of oceanic ecosystems under weak mixing conditions-a theoretical investigation. Prog. Oceanogr. 75:771-96
10. Behrenfeld MJ, Boss E. 2003. The beam attenuation to chlorophyll ratio: an optical index of phytoplankton physiology in the surface ocean? Deep-Sea Res. I 50:1537-49
11. Behrenfeld MJ, Falkowski PG. 1997. A consumer's guide to phytoplankton primary productivity models. Limnol. Oceanogr. 42:1479-91
12. Berdalet E, McManus MA, Ross ON, Burchard H, Chavez FP, et al. 2014. Understanding harmful algae in stratified systems: review of progress and future directions. Deep-Sea Res. II 101:4-20
13. Bidigare RR, Smith RC, Baker KS, Marra J. 1987. Oceanic primary production estimates from measurements of spectral irradiance and pigment concentrations. Glob. Biogeochem. Cycles 1:171-86
14. Bienfang PK, Szyper JP, Laws E. 1983. Sinking rate and pigment responses to light limitation by a marine diatom: implications to dynamics of chlorophyll maximum layers. Oceanol. Acta 6:55-62
15. Birch DA, Young WR, Franks PJS. 2009. Plankton layer profiles as determined by shearing, sinking, and swimming. Limnol. Oceanogr. 54:397-99
16. Boyd CM, Gradmann D. 2002. Impact of osmolytes on buoyancy of marine phytoplankton. Mar. Biol. 141:605-18
17. Briggs N, Perry MJ, Cetinić I, Lee C, D'Asaro E, et al. 2011. High-resolution observations of aggregate flux during a sub-polar North Atlantic spring bloom. Deep-Sea Res. I 58:1031-39
18. Cermeño P, Dutkiewicz S, Harris RP, Follows M, Schofield O, Falkowski PG. 2008. The role of nutricline depth in regulating the ocean carbon cycle. Proc. Natl. Acad. Sci. USA 105:20344-49
19. Cetinić I, Perry MJ, Briggs NT, Kallin E, D'Asaro EA, Lee CM. 2012. Particulate organic carbon and inherent optical properties during 2008 North Atlantic Bloom Experiment. J. Geophys. Res. 117:C06028
20. Chan AT. 1978. Comparative physiological studyofmarine diatoms and dinoflagellatesinrelationtoirradiance and cell size. I. Growth under continuous light. J. Phycol. 14:396-402
21. Claustre H, Huot Y, Obernosterer I, Gentili B, Tailliez D, Lewis M. 2008. Gross community production and metabolic balance in the South Pacific Gyre, using a non intrusive bio-optical method. Biogeosciences 5:463-74
22. Clegg MR, Gaedke U, Boehrer B, Spijkerman E. 2012. Complementary ecophysiological strategies combine to facilitate survival in the hostile conditions of a deep chlorophyll maximum. Oecologia 169:609-22
23. Cowles TJ. 2003. Planktonic layers: physical and biological interactions on the small scale. In Handbook of Scaling Methods in Aquatic Ecology: Measurement, Analysis, Simulation, ed. L Seuront, PG Strutton, pp. 31-49. Boca Raton, FL: CRC
24. Cullen JJ. 1982. The deep chlorophyll maximum: comparing vertical profiles of chlorophyll a. Can. J. Fish. Aquat. Sci. 39:791-803
25. Cullen JJ. 1985. Diel vertical migration by dinoflagellates: roles of carbohydrate metabolism and behavioral flexibility. Contrib. Mar. Sci. 27(Suppl.):135-52
26. Cullen JJ. 2008. Observation and prediction of harmful algal blooms. In Real-Time Coastal Observing Systems for Marine Ecosystem Dynamics and Harmful Algal Blooms: Theory, Instrumentation and Modelling, ed. M Babin, CS Roesler, JJ Cullen, pp. 1-41. Paris: UNESCO
27. Cullen JJ. 2012. Foreword. In Biological Oceanography: An Early History, 1870-1960, by EL Mills, pp. ix-xxi. Toronto: Univ. Toronto Press
28. Cullen JJ, Davis RF, Huot Y. 2012. Spectral model of depth-integrated water column photosynthesis and its inhibition by ultraviolet radiation. Glob. Biogeochem. Cycles 26:GB1011
29. Cullen JJ, Horrigan SG. 1981. Effects of nitrate on the diurnal vertical migration, carbon to nitrogen ratio, and the photosynthetic capacity of the dinoflagellate, Gymnodinium splendens. Mar. Biol. 62:81-89
30. Cullen JJ, MacIntyre HL, Carlson DJ. 1989. Distributions and photosynthesis of phototrophs in sea-surface films. Mar. Ecol. Prog. Ser. 55:271-78
31. Cullen JJ, MacIntyre JG. 1998. Behavior, physiology and the niche of depth-regulating phytoplankton. In Physiological Ecology of Harmful Algal Blooms, ed. DM Anderson, AD Cembella, GM Hallegraeff, pp. 559-80. Berlin: Springer-Verlag
32. Cullen JJ, Reid FMH, Stewart E. 1982. Phytoplankton in the surface and chlorophyll maximum off southern California in August, 1978. J. Plankton Res. 4:665-94
33. Davey MC, Heaney SI. 1989. The control of sub-surface maxima of diatoms in a stratified lake by physical, chemical and biological factors. J. Plankton Res. 11:1185-89
34. Derenbach JB, Astheimer H, Hansen HP, Leach H. 1979. Vertical microscale distribution of phytoplankton in relation to the thermocline. Mar. Ecol. Prog. Ser. 1:187-93
35. Durham WM, Kessler JO, Stocker R. 2009. Disruption of vertical motility by shear triggers formation of thin phytoplankton layers. Science 323:1067-70
36. Durham WM, Stocker R. 2012. Thin phytoplankton layers: characteristics, mechanisms, and consequences. Annu. Rev. Mar. Sci. 4:177-207
37. Eppley RW, Holm-Hansen O, Strickland JDH. 1968. Some observations on the vertical migration of di-noflagellates. J. Phycol. 4:333-40
38. Eppley RW, Peterson BJ. 1979. Particulate organic matter flux and planktonic new production in the deep ocean. Nature 282:677-80
39. Eppley RW, Reid FMH, Cullen JJ, Winant CD, Stewart E. 1984. Subsurface patch of dinoflagellate (Ceratium tripos) off Southern California: patch length, growth rate, associated vertically migrating species. Mar. Biol. 80:207-14
40. Eppley RW, Reid FMH, Strickland JDH. 1970. Estimates of phytoplankton crop size, growth rate, and primary production. See Strickland 1970, pp. 33-42
41. Estrada M, Marrasé C, Latasa M, Berdalet E, Delgado M, Riera T. 1993. Variability of deep chlorophyll maximum characteristics in the Northwestern Mediterranean. Mar. Ecol. Prog. Ser. 92:289-300
42. Falkowski PG, Hopkins TS, Walsh JJ. 1980. An analysis of factors affecting oxygen depletion in the New York Bight. J. Mar. Res. 38:479-506
43. Falkowski PG, LaRoche J. 1991. Acclimation to spectral irradiance in algae. J. Phycol. 27:8-14
44. Fennel K, Boss E. 2003. Subsurface maxima of phytoplankton and chlorophyll: steady-state solutions from a simple model. Limnol. Oceanogr. 48:1521-34
45. Fischer AD, Moberg EA, Alexander H, Brownlee EF, Hunter-Cevera KR, et al. 2014. Sixty years of Sverdrup: a retrospective of progress in the study of phytoplankton blooms. Oceanography 27(1):222-35
46. Fleming RH. 1939. The control of diatom populations by grazing. J. Cons. Int. Explor. Mer 14:210-27
47. Franks PJS. 1992. Sink or swim: accumulation of biomass at fronts. Mar. Ecol. Prog. Ser. 82:1-12
48. Geider RJ. 1992. Quantitative phytoplankton ecophysiology: implications for primary production and phyto-plankton growth. ICES Mar. Sci. Symp. 194:52-62
49. Geider RJ, MacIntyre HL, Kana TM. 1997. Dynamic model of phytoplankton growth and acclimation: responses of the balanced growth rate and the chlorophyll a:carbon ratio to light, nutrient-limitation and temperature. Mar. Ecol. Prog. Ser. 148:187-200
50. Geider RJ, Osborne BA, Raven JA. 1986. Growth, photosynthesis and maintenance metabolic cost in the diatom Phaeodactylum tricornutum at very low light levels. J. Phycol. 22:39-48
51. Gentleman W. 2002. A chronology of plankton dynamics in silico: how computer models have been used to study marine ecosystems. Hydrobiologia 480:69-85
52. Gernez P, Antoine D, Huot Y. 2011. Diel cycles of the particulate beam attenuation coefficient under varying trophic conditions in the northwestern Mediterranean Sea: observations and modeling. Limnol. Oceanogr. 56:17-36
53. Gould WJ. 2005. From Swallow floats to Argo: the development of neutrally buoyant floats. Deep-Sea Res. II 52:529-43
54. Graff JR, Milligan AJ, Behrenfeld MJ. 2012. The measurement of phytoplankton biomass using flow-cytometric sorting and elemental analysis of carbon. Limnol. Oceanogr. Methods 10:910-20
55. Granéli E, Paasche E, Maestrini SY. 1993. Three years after the Chrysochromulina polylepsis bloom in Scandinavian waters in 1988: some conclusions of recent research and monitoring. In Toxic Phytoplankton Blooms in the Sea, ed. TJ Smayda, Y Shimizu, pp. 23-32. Amsterdam: Elsevier
56. Halpern D, Knox RA, Luthier DS, Philander SGH. 1989. Estimates of equatorial upwelling between 140° and 110°W during 1984. J. Geophys. Res. 94:8018-20
57. Hamm CE, Merkel R, Springer O, Jurkojc P, Maier C, et al. 2003. Architecture and material properties of diatom shells provide effective mechanical protection. Nature 421:841-43
58. Hansen G, Daugbjerg N, Henriksen P. 2000. Comparative study of Gymnodinium mikimotoi and Gymnodinium aureolum, comb. nov. (=Gyrodinium aureolum) based on morphology, pigment composition, and molecular data. J. Phycol. 36:394-410
59. Harris G, Heaney S, Talling J. 1979. Physiological and environmental constraints in the ecology of the planktonic dinoflagellate Ceratium hirundinella. Freshw. Biol. 9:413-28
60. Hasle GR. 1950. Phototactic vertical migration in marine dinoflagellates. Oikos 2:162-75
61. Heaney SI, Davey MC, Brooks AS. 1989. Formation of sub-surface maxima of a diatom within a stratified lake and in a laboratory water column. J. Plankton Res. 11:1168-84
62. Heaney SI, Eppley RW. 1981. Light, temperature and nitrogen as interacting factors affecting diel vertical migrations of dinoflagellates in culture. J. Plankton Res. 3:331-44
63. Herbland A, Voituriez B. 1979. Hydrological structure analysis for estimating the primary production in the tropical Atlantic Ocean. J. Mar. Res. 37:87-101
64. IOCCG (Int. Ocean-Col. Coord. Group). 2000. Remote sensing of ocean colour in coastal, and other optically-complex, waters. Rep. No. 3, IOCCG, Dartmouth, Can.
65. Jamart BM, Winter D, Banse K, Anderson G, Lam R. 1977. A theoretical study of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern US coast. Deep-Sea Res. 24:753-73
66. Jamart BM, Winter DF, Banse K. 1979. Sensitivity analysis of a mathematical model of phytoplankton growth and nutrient distribution in the Pacific Ocean off the northwestern US coast. J. Plankton Res. 1:267-90
67. Jenkins WJ, Doney SC. 2003. The subtropical nutrient spiral. Glob. Biogeochem. Cycles 17:1110
68. Jerlov NG. 1959. Maxima in the vertical distribution of particles in the sea. Deep-Sea Res. 5:173-84
69. Ji R, Franks PJS. 2007. Vertical migration of dinoflagellates: model analysis of strategies, growth, and vertical distribution patterns. Mar. Ecol. Prog. Ser. 344:49-61
70. Johnson KS, Berelson WM, Boss ES, Chase Z, Claustre H, et al. 2009. Observing biogeochemical cycles at global scales with profiling floats and gliders: prospects for a global array. Oceanography 22(3):216-25
71. Kamykowski D. 1995. Trajectories of autotrophic marine dinoflagellates. J. Phycol. 31:200-8
72. Karl DM. 1999. A sea of change: biogeochemical variability in the North Pacific Subtropical Gyre. Ecosystems 2:181-214
73. Kiefer DA, Lasker R. 1975. Two blooms of Gymnodinium splendens, an unarmoured dinoflagellate. Fish. Bull. 73:675-78
74. Kiefer DA, Olson RJ, Holm-Hansen O. 1975. Another look at the nitrite and chlorophyll maxima in the central North Pacific. Deep-Sea Res. Oceanogr. Abstr. 23:1199-208
75. Kirk JTO. 2011. Light and Photosynthesis in Aquatic Ecosystems. Cambridge, UK: Cambridge Univ. Press. 3rd ed.
76. Kitchen JC, Zaneveld JRV. 1990. On the noncorrelation of the vertical structure of light scattering and chlorophyll α in case I waters. J. Geophys. Res. 95:20237-46
77. Klausmeier CA, Litchman E. 2001. Algal games: the vertical distribution of phytoplankton in poorly mixed water columns. Limnol. Oceanogr. 46:1998-2007
78. Kononen K, Huttunen M, Hallfors S, Gentien P, Lunven M, et al. 2003. Development of a deep chlorophyll maximum of Heterocapsa triquetra Ehrenb. at the entrance to the Gulf of Finland. Limnol. Oceanogr. 48:594-607
79. Konopka A, Klemer A, Walsby A, Ibelings BW. 1993. Effects of macronutrients upon buoyancy regulation by metalimnetic Oscillatoria agardhii in Deming Lake, Minnesota. J. Plankton Res. 15:1019-34
80. Lande R, Li WKW, Horne EP, Wood AM. 1989. Phytoplankton growth rates estimated from depth profiles of cell concentration and turbulent diffusion. Deep-Sea Res. 36:1141-59
81. Lande R, Wood AM. 1987. Suspension times of particles in the upper ocean. Deep-Sea Res. 34:61-72
82. Langdon C. 1987. On the causes of interspecific differences in the growth-irradiance relationship for phyto-plankton. Part I.A comparative studyofthe growth-irradiance relationshipofthree marine phytoplankton species: Skeletonema costatum, Olisthodiscus luteus and Gonyaulax tamarensis. J. Plankton Res. 9:459-82
83. Lasker R. 1975. Field criteria for survivalofanchovy larvae: the relation between inshore chlorophyll maximum layers and successful first feeding. Fish. Bull. 73:453-62
84. Lee Z, Weidemann A, Kindle J, Arnone R, Carder KL, Davis C. 2007. Euphotic zone depth: its derivation and implication to ocean-color remote sensing. J. Geophys. Res. 112:C03009
85. Letelier RM, Karl DM, Abbott MR, Bidigare RR. 2004. Light driven seasonal patterns of chlorophyll and nitrate in the lower euphotic zone of the North Pacific Subtropical Gyre. Limnol. Oceanogr. 49:508-19
86. Lévy M. 2003. Mesoscale variability of phytoplankton and of new production: impact of the large scale nutrient distribution. J. Geophys. Res. 108:3358
87. Lévy M. 2008. The modulation of biological production by oceanic mesoscale turbulence. In Transport and Mixing in Geophysical Flows, ed. JB Weiss, A Provenzale, pp. 219-61. Berlin: Springer-Verlag
88. Lewis MR, Harrison WG, Oakey NS, Hebert D, Platt T. 1986. Vertical nitrate fluxes in the oligotrophic ocean. Science 234:870-73
89. Li QP, Franks PJS, Landry MR, Goericke R, Taylor AG. 2010. Modeling phytoplankton growth rates and chlorophyll to carbon ratios in California coastal and pelagic ecosystems. J. Geophys. Res. 115:G04003
90. Lorenzen CJ. 1966. A method for the continuous measurement of in vivo chlorophyll concentration. Deep-Sea Res. 13:223-27
91. Lorenzen CJ. 1976. Primary production in the sea. In Ecology of the Seas, ed. DH Cushing, JJ Walsh, pp. 173-85. Oxford, UK: Blackwell Sci.
92. Margalef R. 1978. Life forms of phytoplankton as survival alternatives in an unstable environment. Oceanol. Acta 1:493-509
93. Margalef R, Estrada M, Blasco D. 1979. Functional morphology of organisms involved in red tides, as adapted to decaying turbulence. In Toxic Dinoflagellate Blooms, ed. DL Taylor, HH Seliger, pp. 89-94. New York: Elsevier-North Holland
94. Marra J. 2004. The compensation irradiance for phytoplankton in nature. Geophys. Res. Lett. 31:L06305
95. McGillicuddy DJ Jr, Anderson LA, Bates NR, Bibby T, Buesseler KO, et al. 2007. Eddy/wind interactions stimulate extraordinary mid-ocean plankton blooms. Science 316:1021-26
96. McGillicuddy DJ Jr, Robinson AR. 1997. Eddy-induced nutrient supply and new production in the Sargasso Sea. Deep-Sea Res. I 44:1427-50
97. Michaels AF, Silver MW. 1988. Primary production, sinking fluxes and the microbial food web. Deep-Sea Res. 35:473-90
98. Mignot A, Claustre H, D'Ortenzio F, Xing X, Poteau A, Ras J. 2011. From the shape of the vertical profile of in vivo fluorescence to chlorophyll-a concentration. Biogeosciences 8:2391-406
99. Mignot A, Claustre H, Uitz J, Poteau A, D'Ortenzio F, Xing X. 2014. Understanding the seasonal dynamics of phytoplankton biomass and the deep chlorophyll maximum in oligotrophic environments: a Bio-Argo float investigation. Glob. Biogeochem. Cycles 28:856-76
100. Morel A. 1988. Optical modelling of the upper ocean in relation to its biogenous matter content (Case I waters). J. Geophys. Res. 93:10749-68
101. Morel A, Berthon J-F. 1989. Surface pigments, algal biomass profiles, and potential production of the euphotic layer: relationships reinvestigated in view of remote-sensing applications. Limnol. Oceanogr. 34:1545-62
102. Munk W. 2002. The U.S. Commission on Ocean Policy: Testimony in San Pedro, California, 18
103. April 2002. http://govinfo.library.unt.edu/oceancommission/meetings/apr18-19-02/munk
104. Oliver RL. 1994. Floating and sinking in gas-vacuolate cyanobacteria. J. Phycol. 30:161-73
105. Passow U, Alldredge AL, Logan BE. 1994. The role of particulate carbohydrate exudates in the flocculation of diatom blooms. Deep-Sea Res. I 41:335-57
106. Pingree RD, Pugh PR, Holligan PM, Forster GR. 1975. Summer phytoplankton blooms and red tides along tidal fronts in the approaches to the English Channel. Nature 258:672-77
107. Platt T, Rao DS. 1970. Primary production measurements on a natural plankton bloom. J. Fish. Board Can. 27:887-99
108. Platt T, Sathyendranath S. 1988. Oceanic primary production: estimation by remote sensing at local and regional scales. Science 241:1613-20
109. Platt T, Sathyendranath S. 1993. Estimators of primary production for the interpretation of remotely sensed data on ocean color. J. Geophys. Res. 98:14561-96
110. Prairie JC, Franks PJS, Jaffe JS, Doubell MJ, Yamazaki H. 2011. Physical and biological controls of vertical gradients in phytoplankton. Limnol. Oceanogr. Fluids Environ. 1:75-90
111. Ralston DK, McGillicuddy DJ Jr, Townsend DW. 2007. Asynchronous vertical migration and bimodal distribution of motile phytoplankton. J. Plankton Res. 29:803-21
112. Richardson TL, Ciotti AM, Cullen JJ, Villareal TA. 1996. Physiological and optical properties of Rhizosolenia formosa (Bacillariophyceae) in the context of open-ocean vertical migration. J. Phycol. 32:741-57
113. Richardson TL, Cullen JJ. 1995. Changes in buoyancy and chemical composition during growth of a coastal marine diatom: ecological and biogeochemical consequences. Mar. Ecol. Prog. Ser. 128:77-90
114. Richardson TL, Cullen JJ, Kelley DE, Lewis MR. 1998. Potential contributions of vertically migrating Rhi-zosolenia to nutrient cycling and new production in the open ocean. J. Plankton Res. 20:219-42
115. Riley GA. 1946. Factors controlling phytoplankton populations on Georges Bank. J. Mar. Res. 6:54-73
116. Riley GA, Stommel H, Bumpus DF. 1949. Quantitative Ecology of the Plankton of the Western North Atlantic. Bull. Bingham Oceanogr. Coll. 12. New Haven, CT: Bingham Oceanogr. Lab.
117. Rines JEB, McFarland MN, Donaghay PL, Sullivan JM. 2010. Thin layers and species-specific characterization of the phytoplankton community in Monterey Bay, California, USA. Cont. Shelf Res. 30:66-80
118. Riser SC, Johnson KS. 2008. Net production of oxygen in the subtropical ocean. Nature 451:323-25
119. Roesler CS, Barnard AH. 2014. Optical proxy for phytoplankton biomass in the absence of photophysiology: rethinking the absorption line height. Methods Oceanogr. 7:79-94
120. Rossby H, Levine E, Connors D. 1985. The isopycnal Swallow float-a simple device for tracking water parcels in the ocean. Prog. Oceanogr. 14:511-25
121. Ryan JP, McManus MA, Sullivan JM. 2010. Interacting physical, chemical and biological forcing of phyto-plankton thin-layer variability in Monterey Bay, California. Cont. Shelf Res. 30:7-16
122. Ryther JH. 1956. Photosynthesis in the ocean as a function of light intensity. Limnol. Oceanogr. 1:61-70
123. Sharples J, Moore CM, Rippeth TP, Holligan PM, Hydes DJ, et al. 2001. Phytoplankton distribution and survival in the thermocline. Limnol. Oceanogr. 46:486-96
124. Shulenberger E, Reid JL. 1981. The Pacific shallow oxygen maximum, deep chlorophyll maximum, and primary productivity reconsidered. Deep-Sea Res. 28:901-19
125. Smayda TJ. 1970. The suspension and sinking of phytoplankton in the sea. Oceanogr. Mar. Biol. Annu. Rev. 8:353-414
126. Smayda TJ. 1997. Harmful algal blooms: their ecophysiology and general relevance to phytoplankton blooms in the sea. Limnol. Oceanogr. 42:1137-53
127. Smetacek VS. 1985. Role of sinking in diatom life-history cycles: ecological, evolutionary and geological significance. Mar. Biol. 84:239-51
128. Sommer U. 1982. Vertical niche separation between two closely related planktonic flagellate species (Rhodomonas lens and Rhodomonas minuta v. nannoplanctica). J. Plankton Res. 4:137-42
129. Stacey MT, McManus MA, Steinbuck JV. 2007. Convergences and divergences and thin layer formation and maintenance. Limnol. Oceanogr. 52:1523
130. Steele JH. 1956. Plant production on the Fladen Ground. J. Mar. Biol. Assoc. UK 35:1-33
131. Steele JH. 1962. Environmental control of photosynthesis in the sea. Limnol. Oceanogr. 7:137-50
132. Steele JH. 1964. A study of production in the Gulf of Mexico. J. Mar. Res. 22:211-22
133. Steele JH, Yentsch CS. 1960. The vertical distribution of chlorophyll. J. Mar. Biol. Assoc. UK 39:217-26
134. Steinbuck JV, Genin A, Monismith SG, Koseff JR, Holzman R, Labiosa RG. 2010. Turbulent mixing in fine-scale phytoplankton layers: observations and inferences of layer dynamics. Cont. Shelf Res. 30:442-55
135. Strickland JDH. 1968. A comparison of profiles of nutrient and chlorophyll concentrations taken from discrete depths and by continuous recording. Limnol. Oceanogr. 13:388-91
136. Strickland JDH, ed. 1970. The Ecology of the Plankton off La Jolla, California, in the Period April Through September, 1967. Bull. Scripps Inst. Oceanogr. Vol. 17. Berkeley: Univ. Calif. Press
137. Strom SL, Miller CB, Frost BW. 2000. What sets lower limits to phytoplankton stocks in high-nitrate, low-chlorophyll regions of the open ocean? Mar. Ecol. Prog. Ser. 193:19-31
138. Sullivan JM, Donaghay PL, Rines JEB. 2010. Coastal thin layer dynamics: consequences to biology and optics. Cont. Shelf Res. 30:50-65
139. Taylor AG, Landry MR, Selph KE, Wokuluk JJ. 2014. Temporal and spatial patterns of microbial community biomass and composition in the Southern California Current Ecosystem. Deep-Sea Res. II. In press. doi: 10.1016/j.dsr2.2014.02.006
140. Taylor AH, Geider RJ, Gilbert FJH. 1997. Seasonal and latitudinal dependencies of phytoplankton carbon-to-chlorophyll a ratios: results of a modelling study. Mar. Ecol. Prog. Ser. 152:51-66
141. Taylor AH, Harris JRW, Aiken J. 1986. The interaction of physical and biological processes in a model of the vertical distribution of phytoplankton under stratification. In Marine Interfaces Ecohydrodynamics, ed. JCJ Nihoul, pp. 313-30. Elsevier Oceanogr. Ser. Vol. 42. Amsterdam: Elsevier
142. Tyler MA, Seliger HH. 1978. Annual subsurface transport of a red tide dinoflagellate to its bloom area: water circulation patterns and organism distributions in the Chesapeake Bay. Limnol. Oceanogr. 23:227-46
143. Tyler MA, Seliger HH. 1981. Selection for a red tide organism: physiological responses to the physical environment. Limnol. Oceanogr. 26:310-24
144. Uitz J, Claustre H, Morel A, Hooker SB. 2006. Vertical distribution of phytoplankton communities in open ocean: an assessment based on surface chlorophyll. J. Geophys. Res. 111:C08005
145. Venrick EL. 1988. The vertical distributions of chlorophyll and phytoplankton species in the North Pacific central environment. J. Plankton Res. 10:987-98
146. Venrick EL, McGowan J, Mantyla A. 1973. Deep maxima of photosynthetic chlorophyll in the Pacific Ocean. Fish. Bull. 71:41-52
147. Verity PG, Smetacek V. 1996. Organism life cycles, predation, and the structure of marine pelagic ecosystems. Mar. Ecol. Prog. Ser. 130:277-93
148. - fluxes in the central North Pacific gyre. J. Plankton Res. 18:1103-21
149. Walsby A. 1978. The properties and buoyancy-providing role of gas vacuoles in Trichodesmium Ehrenberg. Br. Phycol. J. 13:103-16
150. Weiler CS, Balch WM, Chisholm SW, Cullen JJ, Harrison WG, et al. 1990. Richard W. Eppley's contributions to phytoplankton physiology and biological oceanography. Oceanography 3(2):42-46
151. White AE, Spitz YH, Letelier RM. 2006. Modeling carbohydrate ballasting by Trichodesmium spp. Mar. Ecol. Prog. Ser. 323:35-45
152. Williams RG, Follows MJ. 2003. Physical transport of nutrients and the maintenance of biological production. In Ocean Biogeochemistry: The Role of the Ocean Carbon Cycle in Global Change, ed. MJR Fasham, pp. 19-51. Berlin: Springer-Verlag
153. Xing X, Claustre H, Blain S, D'Ortenzio F, Antoine D, et al. 2012. Quenching correction for in vivo chlorophyll fluorescence acquired by autonomous platforms: a case study with instrumented elephant seals in the Kerguelen region (Southern Ocean). Limnol. Oceanogr. Methods 10:483-95
154. Xing X, Morel A, Claustre H, Antoine D, D'Ortenzio F, et al. 2011. Combined processing and mutual interpretation of radiometry and fluorimetry from autonomous profiling Bio-Argo floats: chlorophyll a retrieval. J. Geophys. Res. 116:C06020
155. Yentsch CS. 1974. The influence of geostrophy on primary production. Tethys 6:111-18
156. Yentsch CS. 1980. Phytoplankton growth in the sea: a coalescence of disciplines. In Primary Productivity in the Sea, ed. PG Falkowski, pp. 17-31. New York: Plenum