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Primary Productivity Joji Ishizaka (Hydrsopheric, Atmospheric Research Center, Nagoya University)

What is Primary Production?

•Why Primary Production?

•Definition (Biomass and Rate)

•Phytoplankton Groups

•Measurements and Estimation

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生態系ピラミッド

（Saijo, 2002）

/year

/year

/year

/year

Global Carbon Cycle (Sarmiento et al., 2002)

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Characteristics of ocean primary production in the global carbon cycle • Small stock (biomass)

• Large flux (photosynthesis/respiration)

• Large portion of respiration

• No change after human activity ?

Definition of Primary Production

•Rate at which energy is stored by photosynthetic or chemosynthetic action of producer organisms (autotrophic organisms), in the formation of organic substances

• Production of basic organic materials for heterotrophic organisms

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Primary Producers in the Ocean

•Phytoplankton

•Macroalgae

•Attached microalgae

•Chemosynthetic Bacteria

Photosynthesis

6CO2 + 12H2O +Light Energy → 6CH2O + 6H2O + 6O2

Gross Primary Production (Photosynthesis) Net Primary Production (Photosynthesis - Respiration)

Respiration 6CH2O + 6H2O + 6O2 → 6CO2 + 12H2O + Energy

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Chemosynthesis (sulfur bacteria)

H2S + 2O2 → H2SO4 + Energy

6CO2 + 6H2O +Energy

→ 6CH2O + 6O2

Biomass and Production Biomass (Concentration)

Mass/Volume : g-chl.a/L, = mg-chl.a/m3

Mass/Area : mg/m2

Production (Rate)

Mass/Volume/Time : mgC/L/day

Mass/Area/Time : mgC/m2/day, gC/m2/year

Production/Biomass (Activity) PB

Mass/Mass/Time : mgC/mg-chl.a/day

Function of Temperature, Light, Nutrients, Species...

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Phytoplankton Groups • Cyanobacteria (Blue-Green Algae) • Rhodophyceae (Red Algae) • Cryptophyceae (Crypotomonads) • Chrysophyceae (Chrysomonads, Silicoflagellates) • Bacillariophyceae (Diatoms) • Raphydophyceae (Chloromonads) • Xanthophyceae (Yellow-Green Algae) • Eustigmatophyceae • Prymnesiophyceae (Coccolithophorids, Prymnesiomonads)• Euglenophyceae (Euglenoids) • Prasinophyceae (Prasinomonads) • Chlorophyceae (Green Algae) • Dinophyceae (Dinoflagellates)

Cyanobacteria Trichodesmium Synechococcus Prochlorococcus

Red tide in tropical waters, N2 fixation

Recently found Abundant in Open Ocean

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Diatoms (Centric, Pennate)

Chaetoceros Navicula

Most Abundant in Coastal Water Shell of Silica

Diatoms

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Coccolithophorids

Plates of CaCO3 Sometimes form white water

Coccolithophore Bloom in Araska 1997 Spring

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Dinoflagellates

Ceratium Gimnodinium

Some times forms Harmful Algal Bloom or Red Tide Some species are heterotrophic

Dinoflagellate (HAB species)

(Raphidophyte)

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Food Chain

Lalli and Parsons

Zooplankton （Protozoa）

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Zooplankton（Copepods）

Ryther, 1969 (Science)

Area

(x106

km2)

Primary

Production

(gC/m2/y)

Number

of Trophic

Level

Transfer

Efficient

(%)

Fish

Production

(mgC/m2/y)

Global Fish

Production

(x103 t/y)

Open

Ocean 322 50

(1)

5 10 0.5

(1)

150

(1)

Coastal 26.2 100

(2)

3 15 340

(680)

9000

(60)

Upwell-

ing 0.4 300

(6)

1.5 20 36,000

(72,000)

14,400

(100)

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Estimation of Global Primary Production

Riley (‘46) 126 DO method, several stations

Steemann Nielsen (‘55) 15 14C methods

Ryther (‘69) 20 Ocean:Coast:Upwelling= 90:9.9:0.1

Koblentz-Mishke et al.(‘70)

23 7000 data

Lieth and Whittaker (‘75) 18.6 Fleming (‘57)

Platt and Sabbarao (’75) 31 Summaries areal data

Eppley and 19.1 Modified Koblentz-Mishke et al. (‘70)

Peterson (‘79) 23.7 Modified Platt and Subba Rao (‘75)

Romankevich(‘84) 25 Modified Koblentz-Mishke et al. (‘70)

Shushkina (‘85) 56 130 stations (‘68-’82)

Berger et al. (‘87) 26.9 8000 stations (mostly ‘70-)

Martin et al. (‘87) 51 Ryhter (‘69) method + Clean Method

Longhurst et al. (‘95) 45-51

Satellite data (CZCS) + biological provinces

Measurements of Primary Production by 14C/13C

Light Bottle

Dark Bottle

Net Primary Production

Blank

+NaH1４CO3

Organic 14C・13C

Sampled water in bottle Add 14C or 13C（HCO3

- ） hrs～１day incubation （Same Light Condition） Carbon in Organic Matter

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Incubation In situ Method

Simulate In situ Method

Sverdrup (1955) The Place of

Physical Oceanography in the

Oceanographic Research

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Longhurst et al. (1995)

Controlling Factors • Phytoplankton Biomass

• Light (Energy Source)

• Nutrients (Material Source)

–Macro: NO3, NO2, NH4, PO4, SiO2

–Micro: Fe,,,

• Temperature (Chemical Reaction Speed)

• Phytoplankton Group

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Environmental Control of Photosynthesis

TemperatureNutrientsLight

PB

Light is required for energy.

Nutrients are required for material.

Temperature controls rate of chemical reaction.

Satellite Primary Production Model

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Chl-a, SST, PAR → GLI Primary Production (April-June 2004)

JAXA

Different Type of Primary Production Modelｓ (Behrenfeld and Falkowski, 1997)

• Wavelength-Resolved Models (WRMs) -(WTDRM)

• Wavelength-Integrated Models (WIMs)-(TDRM)

• Time Integrated Models (TIMs)-(DRM)

• Depth-Integrated Models (DIMs)-(IM)

700

400 0

* )(),,(),(

Sunset

Sunrise

Zeu

Z

dtdzdzChlaztPARzPP

Sunset

Sunrise

Zeu

Z

dtdzzChlztPARzPP0

)(),()(

Zeu

Z

B dzzChlDLzPARzPPP0

)()()(

ZeuChlDLPARPPP optB )0()0(

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Depth Resolved Primary Production Model

PB

PB

Wavelength, Time, Depth Resolved Primary Production Model (Ishizaka et al., 1998) based on (Morel, 1991)

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Off Sanriku Apr. 26, 1997 Oyashio, Warm Core Ring, Kuroshio

OCTS derived Primary Production off Sanriku

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Chl.a(0)

Day Length (DL) P

AR

Euphotic

Zone

(Zeu)

Integrated

Chl.a

Surface

Zeu

Dep

th

PP/Chl.a

PBopt

IPP

VGPM (Vertically Generalized Production Model) Behrenfeld & Falkowski (1997)

eu

b

opt ZaChlDLPARfPIPP 0.0

IPP : Integrated Primary

Production

・Behrenfeld&Falkowski(1997)

→Pbopt (7th Order Polynomial

of SST)

Modified Behrenfeld and Falkowski (1997)

Two Phytoplankton Community Model of PBopt

(Kameda and Ishizaka, 2005)

IPP =

0.66125 PBopt [E0 / (E0 + 4.1)] Zeu Cz0 Dirr

PBopt = f (SST) B&F 7th order polynomial

PBopt = f (SST, CHL) K&I

CHL = CHLS + CHLL

Prod = ProdS CHLS + ProdLCHLL

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Temperature Dependence of PBopt

(Behrenfeld and Falkowski, 1997)

Temperature-Chlorophyll Dependent Model of PB

opt (Kameda & Ishizaka, 2005) PB

opt = (0.071 T -3.2E-3 x T2 + 3.0E-5 x T3)/Chl

+ (1.0 + 0.17 T – 2.5E-5 x T2 – 8.0E-5 x T3)

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Estimating marine primary productivity in coastal and

pelagic regions across the globe: An evaluation of satellite-based

ocean color models Biogeoscience Discussion

Vincent S. Saba, Marjorie A. M. Friedrichs, David Antoine, Robert A.

Armstrong, Ichio Asanuma, Michael J. Behrenfeld, Aurea M. Ciotti, Mark

Dowell, Nicolas Hoepffner, Kimberly J. W. Hyde, Joji Ishizaka, Takahiko

Kameda, John Marra, Frédéric Mélin, André Morel, John O’Reilly, Michele Scardi, Walker O. Smith Jr., Tim J. Smyth, Shilin Tang, Julia Uitz, Kirk

Waters, Toby K. Westberry

(Yamada et al., 2006-JO)

Primary Production in the Japan Sea by K&I Model (Yamada et al, 2005)

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Verification of K&I Model in the Sea of Japan

↑with in situ PBopt

↓Verification of PBopt

（Yamada et al, JO-2005)

(Siswanto et al., 2006-JO)

Primary Production Measurements in the East China Sea

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(Siswanto et al., 2006-JO)

B&F, K&I In situ PBopt

Verification of VGPM in the East China Sea

B&F

K&I

Depth Dependent Model

VGPM

Siswanto

(East ECS）

Gong & Liu

(West ECS）

K&I

B&F

Verification of PBopt

in the East China Sea

(Siswanto et al., 2006-JO)

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● 02

●

●

●

●

● F1

S1 S2

Sagami Bay

1999-2003

(49 Stations）

Tokyo Bay

(Ishizaka et al.,

JO-2007)

Verification in the Sagami Bay Behrenfeld & Falkowski (1997) Kameda & Ishizaka

Integrated

Primary

Production

PBopt

(Ishizaka et al.,

JO-2007)

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PBopt and

SST , Chl(0), PAR

SST

Chl(0)

PAR

(Ishizaka et al., JO-2007)

PBoptModel in the Sagami Bay

PBopt =

(0.071 T -3.2E-3 + 3.0E-5 T3)/Chl

+ (1.0 + 0.17 T – 2.5E-5 T2 – 8.0E-5 T3)

PBopt =

(-12.2+1.17 T - 0.025 T2)/Chl

+ (13.3 + 0.916 T – 0.0191 T2)

(Ishizaka et al., JO-2007)

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Predicted PBopt

B&F

K&I

Sagami Bay Model

Parameter Adjustment of VGPM

(original B&F) 0.66125 PBopt PAR/(PAR+4.1) DL Chl(0) Zeu

In situ PBopt

(best fit) 4.19 PBopt PAR/(PAR+336)DL Chl(0) Zeu

In situ PBopt

IPP=

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Results of Sagami Bay Model and Global Models

RMSE=247

RMSE=392

RMSE=1182

Sagami Bay Model

B&F

K&I

ARIAKE BAY

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(a)modeled PB

opt and Zeu (b) in situ PBopt and Zeu

(c) modeled PBopt and in situ Zeu (d) in situ PB

opt and modeled Zeu With in situ Chl-a

• (a) BF suggested and in situ PB

opt

• (b) MB suggested and in situ Zeu

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(a) in situ 1/Zeu and anph (b) calculated Zeu from multiple regression relationship and measured Zeu (c) QAA-based Zeu and in situ Zeu.

● PBopt variations based on datasets of this study,

○ model of Behrenfeld and Falkowski (1997), ◇ SST-dependent model of model of Gong and Liu (2003), △ model of Kameda and Ishizaka (2005) □ model of Siswanto et al. (2006)

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(a)original OC4v4 (b) optimized OC4v4 (c) original Tassan (d) optimized Tassan

(a)13C-based in situ IPP (b) FRRF single instantaneous cast-based

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• Japan Sea: OK with open ocean model.

• East China Sea: Pbopt SST dependency

• Sagami Bay: Pbopt, Integration(?)

• Ariake Bay (Turbid): Pbopt, Zeu, Chl-a

Summary • Primary Production is important parameter

for ecosystem dynamics.

• With Satellite, it is easy to calculate.

• Some models with different complexity are available.

• However, parameters are different regionally, and at present measurements and tuning of the model is necessary, especially coastal environment is challenging.

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References and suggested readings • Ishizaka, J., E. Siswanto, T. Itoh, H. Murakami, Y. Yamaguchi, T. Ishimaru, S.

Hashimoto and T. Saino (2007): Verification of Vertically Generalized Production Model and Estimation of Primary Production in the Sagami Bay, Japan. J. Oceanogr. 64, 5１7-524.

• Siswanto, E., J. Ishizaka and K. Yokouchi (2006) Optimal primary production model and parameterization in the eastern East China Sea. J. Oceanogr. 62, 361-372.

• Yamada, K., J. Ishizaka, H. Nagata (2005) Spatial and temporal variability of satellite estimated primary production in the Japan Sea from 1998 to 2002. J. Oceanogr. 61, 857-869.

• Kameda, T. and J. Ishizaka (2005) Size-fractionated primary production estimated by a two-phytoplankton community model applicable to ocean color remote sensing. J. Oceanogr. 61, 663-672.

• Carr, M. E., M.A.M. Fredrichs, M. Schmeltz, M.N. Aita, D. Antoine, K.R. Arrigo, I. Asanuma, O. Aumont, R. Barber, M. Behrefeld, R. Bidigare, E. Buitenhuis, J. Campbell, A. Ciotti, H. Dierssen, M. Dowell, J. Dunne, W. Esaias, B. Gentili, S. Groom, N. Hoepffner, J. Ishizaka, T. Kameda, C. LeQuere, S. Lohrenz, J. Marra, F. Melin, K. Moore, A. Morel, T. Reddy, J. Ryan, M. Scardi, T. Smyth, K. Turpie, G. Tilstone, K. Waters, Y. Yamanaka (2006): A compariosn of global estimates of marine primary production from ocean color. Deep-Sea Res. II, 53, 741-770.

References and Suggested Readings

• Ishizaka, J. (1998): Spatial distribution of primary production off Sanriku, Northwestern Pacific, during spring estimated by Ocean Color and Temperature Scanner (OCTS). J. Oceanogr., 54, 553-564.

• Behrenfeld, M. J. and P. G. Falkowski (1997): A comumers guide to primary productivity models. Limnol. Oceanogr. 42: 1479-1491.

• Behrenfeld, M. J. and P. G. Falkowski (1997): Photosynthetic rates derived from satellite-based chlorophyll concentration. Limnol. Oceanogr. 42: 1-20.

• Longhurst, A., S. Sathyendranath, T. Platt and C. Caverhill (1995): An estimate of global primary production in the ocean from satellite radiometer data. J. Plankton Res. 17: 1245-1271.

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References and Suggested Readings • 2010 Saba, V.S., M.A.M. Friedrichs, M.-E. Carr, D. Antoine, R.A. Armstrong,

I. Asanuma, O. Aumont, N.R. Bates, M.J. Behrenfeld, V. Bennington, L. Bopp, J. Bruggeman, E.T. Buitenhuis, M.J. Church, A.M. Ciotti, S.C. Doney, M. Dowell, J. Dunne, S. Dutkiewicz, W. Gregg, N. Hoepffner, K.J.W. Hyde, J. Ishizaka, T. Kameda, D.M. Karl, I. Lima, M.W. Lomas, J. Marra, G.A. McKinley, F. Melin, J.K. Moore, A. Morel, J. O’Reilly, B. Salihoglu, M. Scardi, T.J. Smyth, S. Tang, J. Tjiputra, J. Uitz, M. Vichi, K. Waters, T.K. Westberry, and A. Yool, Challenges of modeling depth-integrated marine primary productivity over multiple decades: A case study at BATS and HOT, Global Biogeochem. Cycles, 24, GB3020, doi:10.1029/2009GB003655.

• Saba, V.S., M.A.M. Friedrichs, D. Antoine, R.A. Armstrong, I. Asanuma, M.J. Behrenfeld, A.M. Ciotti, M. Dowell, N. Hoepffner, K.J.W. Hyde, J. Ishizaka, T. Kameda, J. Marra, F. Melin, A. Morel, J. O'Reilly, M. Scardi, W.O. Smith Jr., T.J. Smyth, S. Tang, J. Uitz16, K. Waters, and T.K. Westberry (2011) An evaluation of ocean color model estimates of marine primary productivity in coastal and pelagic regions across the globe, Biogeosciences, 8, 489-503.