C-MORE 2012: Measuring phytoplankton productivity & biomass Oscar Schofield

Scalar visible irradiance

Photosynthesis = PAR *

*f

Quantum efficiency of ..

Absorption

Spectrally averaged absorption

700nm
Irradiance 0.00
0.00 0.20 0.40 0.60 0.80 1.00

a ph = 400nm

a(l ) * Eo(l )dl

700nm 400nm

Fluorescence or charge separation oxygen

Eo(l )dl

20.00 D ep t h ( m )

40.00

Energy that is into the cell Varies with cell pigmentation, light history, and size
dEd ( ) K d ( ) E d ( ) dz
~ 1%

60.00

carbon fixation Low growth

80.00

100.00

Energy into the ocean Varies with depth according to the IOPs IOPs with radiative transfer eqns describe the AOPS

Efficiency of converting energy into End product (electrons, oxygen, carb Varies with end product and physiolo

High

Phytoplankton growth and nutrient assimilation is tied to ambient light levels.

Enough energy to excite (vibrate a molecule)

Enough energy move electrons

Enough energy make something new (rearrange a molecule)

Every day, the ocean changes colour or rather, it passes though a variety of hues between the morning, noon and night of a single day. The subtle shapes of clouds, the glittering light of the sun, and the shifts in atmospheric pressure tint the sea with deep tones, cheerful tomes, plaintive tones that would cause any painter to pause in wonder. from The Samurai by Shusaku Endo (1980)

Eyeball Optics
The Secchi Disk:

First systematic usage reported in 1866, but observed and remarked upon much earlier.
Early experiments carried out by Commander Cialdi, head of the Papal Navy, and Professor Secchi onboard the SS LImmacolata Concezione (Cialdi, 1866).

Used operationally for establishing aids to navigation over shallow water.

Thanks to Marlon Lewis

Eyeballs Watch Harmful Algal Blooms HABs

Optical Properties of the Sea

The underwater light field is not a collimated beam. So we define another term(s), the diffuse attenuation coefficient(s), K, to describe the penetration of light in the sea. It is closely related to absorption. Irradiance
0.00 0.20 0.40 0.60 0.80 1.00 0.00 20.00

Depth (m)

40.00 60.00 80.00 100.00

dEd ( ) K d ( ) E d ( ) dz

~ 1%

Measuring the light into the system

From John T. O. Kirks billabongs

Reflectance() = G* bb()/{bb() +a()}

primary productivity

(Behrenfeld and Falkowski, 1997)

or export productivity

(Muller-Karger et al., 2005)

What are some of the classical DIMS?

Y* = P/(Qpar(0+)<Chl>)
Morel and Platt show Y* variability Of 50% around a value of at specific chl values

Local weather

seasonal

Claustre et al.

Diatoms Y 0.114 + 0.051 Cryptophytes Y 0.053 + 0.011

Penetration of light is determined by the material in the water which is determined by the overall inherent optical properties (IOPs)

Absorption (a) color

Photos by S. Etheridge

 Scattering (b) clarity a+b=c c = attenuation

From Collin Roesler

WetLabs

detector

detector

Absorption (a) Attenuation (b)

Scattering Case I waters

cp(660) bp(660)

c p (660 ) A Chl B

Positively correlated Non-linear, B 0.7 High unexplained variance

Loisel and Morel 1998

Particle backscattering
West Florida Shelf

Karenia brevis bloom

Cannizzaro et al. 2002

POC Scattering (Case I waters)

Subtropical Pacific, North Atlantic

c p (660 ) A POC B
B1 Contrast with non-linear dependence on Chl POC-Chl variations are important

Loisel and Morel 1998

POC-Scattering
Bottom layer
New England Continental Shelf

cp(660)

Surface layer

Gardner et al. 2001

Under specific conditions, the cp POC relationship varies

From beautiful work of Morel, Bricaud, and Kirk

E.m. radiation propagating as plane waves; g geometric cross section (its shadow )

EFFICIENCY FACTORS
Energy absorbed within Energy scattered out by.. Divided by Energy impinging on g

Qa and Qb, respectively

(Finkel 2001)

Durand et al. 2002

Positively correlated Chl: 0.02 25 mg m-3

(eutrophic, mesotrophic, and oligotrophic waters) Non-linear dependence
Thanks to Heidi Sosik

Bricaud et al. 1995

 variability in aph()
a ( )
* ph

Chl-specific phytoplankton absorption Second-order

a ph ( ) Chl

Negatively correlated

a * ( ) A( ) Chl B ( ) ph
A() and B() statistically determined

This reflects effects of changing growth conditions and community structure with trophic status

Note: unexplained variability

High light

Low light

Photosynthetic pigments

Low light

Photo-protective pigments

High light

chlorophyte alga Haematococcus pluvialis

chl a
1.25

chl c chl b

phycobilins carotenoids

chl b chl a Spectral Irradiance (

chl a-chl c-carotenoids 20 chl a-chl b-carotenoids chl a-phycobilins

15

Relative Absorption

1.0
0.75

10 0.50 5 0.25 0 400 450 500 550 600 650 700 0

mW cm-2 nm-1)

Wavelength (nm)

mol photons (m-2 s-1)

0 2000 2000 2 6 8

1.2 1.2 1.2 0.9 0.9 0.9 0.6 0.6 0.6 0.3 0.3 0.3
surface 1m

1500 1500 4
h

1000 1000
t

2m
5m 13m

1 500 0 500 1 2
p
m ) ( ( ( m m D ) )e

0 20 0
0 0 0 2 2 2 4 4 4 6 6 6 8 8 8 0 0

0 J

4 2

6 2

8 2

1 2 1 0 D Day a 0

2 y

u lCalendar i a n 0 5

Depth (m)

1 1 1 1

1 8 0 8
0 2

2 0 2 1 2 0 2 2 0

2 2

0 4 0 0 2 0 4 2 J 2 l 0 4 0 J u l u

6 2 6 2 i 6 2 i

0 8 1 2 2 0 0 8 1 2 2 0 a n D 0 8 1 2 2 0 a n D

1 2 1 1 2 1 a y 1 2 1 a y

4 4

1 2 1 0

h D De e p p p t t h h D e t

D e e t D D ep pp t th hh

Depth (m) ((

01 00 1 5 0 0 0

m (m m ) )

0 C 0 0 400 400 400

450 450 450

500 550 600 650 700 500 550 600 650 700 500 550 600 Wavelength (nm) 650 700

2 0
2 4 4 4 6 6 6 8

D2 2
2

4 2

4 2

0 l

6 2 i

6 2

8 2

8 2

0 2

0 2

2 y

n 0J 4u 0l 6i 0 8 1 2 2 a 2
J u a n

u l Calendar i a n
0 6 0 6

Day D

D a

D 0 1 2 1 2 a y

u l Calendar i a n

Day D

Oliver et al. 2004

31 32 1 9 0 0 6 7 31 32 1 9 0 0 6 7 0 6 photons 9 -2 s-1 6 7 mol 3 1 3 2 1 m 0 0

0 0 0 0 0 . . 0 0 .0 0 1 .6 0 3 .1 4 7 0 .0 1 .6 3 .1 4 7

. 0 .a 0 .6 -1 .1 4 0 0 1 (m 03)
ph

Individual pigments can be measured on discrete samples biochemically

absorption coefficient (m2 mg-1)

0.08 0.06 0.04 0.02

chl a chl b chl c PSC PPC

0.0 400 450 500 550 600 650 700

wavelength (nm)
From Bidigare

Spectral irradiance (mWcm-2s-1)

10 1 1 0.1 0.01 0.001 90

B)

25

Depth (m)

Sun stimulated fluorescence

20 40 60 80

Chl a Chl b PSC Chl c

0.0001

A)
500 600 700 100
Decreasing efficiency Increasing efficiency

0.00001 400

Wavelength (nm)

Relative pigment-specific spectrally weighted absorption

Heat

Absorbed photon

Charge stabilization & photosynthesis

Fluorescence

Chlorophyll a - Fate of photons absorbed by an isolated molecule

Diagram of energy states in chlorophyll and possible transitions

Light Absorption

Heating

Thermal Expansion Alexander Graham Bell developed spectrophone, essentially an ordinary spectroscope equipped with a hearing tube instead of an eyepiece listening to light induced changes in the thermal sound.

Pressure Wave

Photoacoustic signal

Strong light flash

Weak light flash

Heat

Absorbed photon

Charge stabilization & photosynthesis

Fluorescence

Fluorescence

Use of sun-induced chlorophyll fluorescence to estimate the rate of carbon fixation -Example

Pacific

Stegmann et al. (1992) JGR

Chlorophyll fluorescence

Stress (light, nutrients)

Chlorophyll concentration

monospecific G. breve community

0
5

0 5

Depth (m)

10 15 20

Depth (m)

10

15
20

mixed chromophyte community

CDOM
8:00 12:00 18:00 22:00

Chl a 8:00 12:00 18:00

22:00

Local DaylightTime

Local DaylightTime

Physiological response

Environmental Stress
PAR UVA + PAR UVB + UVA + PAR

0.75

1600

0.6

Visible light downregulation

800

EPS

1200 0.50 800 0.25

PAR (mmol m-2 s-1)

PAR (mmol m-2 s-1)

600 0.4

Fv/Fm

UVB damage

400

Fv/Fm

0.2 200
Ik>PAR Ik>PAR

400

0.00
5:00 9:00 13:00 17:00

0
21:00

0 6:00 10:00 14:00 18:00 Local Daylight Time

Local Daylight Time

Falkowski et al. (1991) Nature

Fluorescence: The Basics

Fm Fluorescence intensity Fm

F0 = a ph PAR

kf k p + k f + kd kf k f + kd kf k p (Q)+ k f + kd

Fm = a ph PAR
Ft

Fv = a ph PAR

F0

Saturating flash

kp Fm - F0 o = = f IIe Fm k p + k f + kd

time

Integrated area is reflection of the absorption cross-section

Other useful indices FLUORESCENCE INDUCTION

Photons

Low light cells Fluorescence rise

High light cells

Photo-acclimation

Time Flash is on

RC2

Other useful indices FLOURESCENCE DECAY CONSTANTS

Fluorescence

Fluorescence decay constants

Light Flash Turned Off Time D2 Qa Pheo RC D1

0.7

0.6

0.5

0.4

0.3

PQ

0.2

Qb

0.1

0 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800

Local time of day

Heat

Absorbed photon

Charge stabilization & photosynthesis

Fluorescence

Light Reactions produce ATP, NADPH2

Transmission & light absorption

ATP & NADPH

Light

Light Reactions
CO2

Dark Reactions
sugars & carbos Nitrogen, Phosphorus, Metals

Biomass Increase

Cellular Growth

Carbon Quantum Yield (mol C mol photons absorbed-1)

0.1

10

0.08

Productivity (mg C mg Chl a-2 h-1)

fmax

0.8

8
0.7

0.06

Fv/Fm

Pmax
Environmental stress

0.6

0.04

0.02

a
2

0.5

0 0.1 1 10 100

Ik

0.4

Irradiance (mmol photons

1000 m-2 s-1)

Production = Pmax . tanh(PAR/Ik)

4

oxygen evolution

3 2 1 0

Pmax

a
Ik = Pmax/a

50

100

150

200

250

300

PAR (mmol photons m-2 s-1)

Predicting oxygen evolution from fluorescence (model assuming FII = 0.5)

measured oxygen evolution
6 5 4 3

2 1

R2=0.92, P<<0.0001
0 0 1 2 3 4 5 6

fluorescence-based predictions of oxygen evolution

Is a cell a puddle or lake?

From Jassby and Platt 1976

a (mg C mg chl a-1 h-1[mmol photons m-2 s-1]-1)

0.02 0 20 0.03 0.04 0.05 0.06 0.07 0.08

Chl-specific alpha

light-saturated

depth (m)

40 60 80 100 light-limited

aw

Photosynthesis a chain of cascading reactions: Each step sets the upper limit efficiency for each following step down the line Fmpsii (0.65) > fm02 (on the order 0.125)>fmco2 (on the order 0.07) For each use of energy go to one process, it is the expense of another reaction, this impacts the overall efficiency Nutrient source Ammonium Nitrate fmco2 0.09 0.07

Simplest expression for photosynthesis is

P = f * aph * PAR

From Behrenfeld et al.

The conversion efficiency can varies between end products

Ratio of Oxygen to Carbon Quantum Yields Light-limited photosynthesis
12 10 8 6 4 2 0 0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 5

Light-saturated photosynthesis

Absorbed Quanta by phytoplankton

While chlorophyll specific absorption varies 3-4 fold quantum yields vary by an order of magnitude
Even in 1980s was treated as a constant

From Babin et al.

NUTRIENT LIMITATIONS
Iron-Ex

Rough seas

fmax (mol C mol photons absorbed-1)

0.125 0.105 0.085 0.065 0.045

11 8 9

10

4 5 6

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

Arabian Sea (SW Monsoon) Arabian Sea (Inter Monsoon) Arabian Sea (NE Monsoon) Antarctic (Bransfield-Bellingshausen water) Antarctic (Bransfield-Weddell water) Antarctic (Ice-Edge water) Antarctic (Weddell-Scotia Confluence waters) Antarctic (Bellingshausen Cold water) Antarctic (Bellingshausen Warm water) Antarctic (Transitional Weddell Water) Antarctic (Palmer Station) Canary Islands (Fall) NE Atlantic Subtropical Gyre (Fall) NW Atlantic Subtropical Gyre (Fall) Gulf Stream (Fall) NW Atlantic Continental Shelf (Fall) Canary Islands (Spring) NE Atlantic Subtropical Gyre (Spring) NW Atlantic Subtropical Gyre (Spring) Gulf Stream (Spring) NW Atlantic Continental Shelf (Spring) Southern California Bight New Jersey Coastal Region

0.025
21

23 13 15 7 16 14

22

20 19 18 17

1 2

12

Oligotrohic seas
250 300 350

0.005

50

100

150

200

Ek(PAR) (mol photons m-2 s-1)

Types of satellite models

Empirical: Purely a statistical fit. Earliest ones were simple regression models of measured primary production against satellite derived chlorophyll concentration. These models can explain a lot of the observed variance; however assume that the fraction of productivity per phytoplankton cell is essentially fixed.
Ck = surface chlorophyll concentration, II or Pt = depth-integrated production
Smith & Baker 1978

Eppley et al. 1978

Early models were derived by Smith et al. (1982) and Eppley (1985) at Scripps Visibility Labs and Food Chain Working Group

Types of satellite models

Empirical: It works because, the amount of chlorophyll in a volume of water is proportional to the number of phytoplankton cells and the number of photosynthetic reaction centers. In many ways it provides a good Ockams razor for the sophisticated productivity models.
Applying the razor: Back in early 1990s, many sophisticated bio-optical productivity were being developed. A global comparison of the models at the time indicated that simple correlation models did as well or even sometimes better than more sophisticated and biologically realistic models. This did not sit well with some in the community.

Complexity can add uncertainty

Problems: l These relationships assume that the conversion of light energy is constant (not true), that the light harvested for within and between phytoplankton is constant (not true). l Chlorophyll is a concentration, productivity is a rate which has a time dependent variable. l Scales in which to apply the model?

The overcome the errors associated with the depth-dependent variability You generate climatalogies and idealized profiles.

Ignoring the vertical behavior can lead to a 30% error

Remember the majority of phytoplankton biomass is likely light limited, so the importance of the upper water column where light levels are high dominate the integrated productivity estimates.

Platt and Sathyendranath, Science

Relative cloud cover

Biogeographic provinces

Monthly weighted chl

productivity

Platt and Sathyendranath, Science