net primary production (npp) = f [biomass, physiology, g (i )] · 2008. 11. 10. · net primary...
TRANSCRIPT
-
Net primary production (NPP) = f [biomass, physiology, g (I0)]
NPP = f [chlorophyll, Zeu, Pbopt, g (I0)]
{0.02 to 30 mg m-3}
{120 m}
{0 to 30 mg C mgChl-1 h-1}
{0 to 1}
{2 to 300 mg m-2}
NPP = f [Carbon, Zeu, , g (I0)]
{5 to 100 mg m-3} {0 to 2 div. d-1}
-
Light scattering # particles
Common measures of scatter: particulate beam attenuation (cp)particulate backscatter coeff. (bbp)
Open—ocean particle size spectra relatively conserved
cp is:
dominated by particles in the phytoplankton size domain
sensitive to inorganic particles in the 0.5 to 20 m size range
insensitive to submicron bacteria, ‘pure water’ values are measured when 106 bacteria are present
(1) (2)
(3)
(4)
10-1
100
101
102
0
0.05
0.1
0.15
0.2
0.25
0.3
0.35
0.4
0.45
s ize [µm]
beam
atte
nuat
ion
/ pa
rticl
e vo
lum
e
Detrital-like (n=1.04+i*0.0001)
P hytoplankton-like (n=1.05+i*0.005)
10-1 101 102100
0.4
0.3
0.2
0.1
0
beam
atte
nuat
ion
/ par
ticle
vol
ume
size (um)
detrital particles
planktonic particles
14 Steps
-
Particle # cp per particle
cp covaries with POC
Bishop, J.K.B. 1999. Deep-Sea Res. I. 46:353-369.Bishop, J. K. B., S. E. Calvert and M. Y. S. Soon (1999) Deep-Sea Res. II 46:2699-2733.Claustre, H., A. Morel, M. Babin, C. Cailliau, D. Marie, J-C Marty, D. Tailliez and D. Vaulot (1999) J. Geophys. Res. 104:3401-3422.Gardner, W. D., I. D. Walsh and M. J. Richardson (1993) Deep-Sea Res. II. 40:171-195.Gardner, W. D., S. P. Chung, M. J. Richardson and I. D. Walsh (1995) Deep-Sea Res. II. 42:757-775.Loisel, H. and A. Morel (1998) Limnol. Oceanogr. 43: 847-858.Walsh, I. D., S. P. Chung, M. J. Richardson and W. D. Gardner (1995) Deep-Sea Res. II 42:465-477.
Therefore – cp should be a better estimate of phytoplankton carbon than POC
If true – then the ratio of cp:chlorophyll should track phytoplankton C:Chl,which is a common index of physiological variability that sharesenvironmental dependencies with Pbopt and Ek
(5)
(6)
(7)
(8)
0.1 1 10 100 1000 m
Practical POC
Particulate beam attenuation (cp)
-
cp:chl tracks changes in Pbopt and Ek
Behrenfeld & Boss (2004) Deep-Sea Res.50:1537-1549
(9)
0 10 20 30 40 50
0
3
6
9
12
15
18
0.0
0.3
0.6
0.9
1.2
1.5
1.8
0 10 20 30 40 50 60 70
0
3
6
9
12
15
18
0.0
0.3
0.6
0.9
1.2
1.5
1.81992 1993 1994 1995 1996 1997
* *
1994199319921991 19961995
April May June
0
2
4
6
8
0.0
0.2
0.4
0.6
0.8
1.0
653 4130 2 272523201886 312927 292625
NABE Time Series
Sequential Observation
Mea
sure
d Pb
opt
Measured c
p :Chl ratio
NABE
BATS
HOT
Mea
sure
d Pb
opt
cp :C
hl ratio
0.0 0.2 0.4 0.6 0.8 1.00
100
200
300 OliPac
cp:Chl ratioM
easu
red
E k
-
Ocean color inversion algorithms give us bbp , not cp
Mie calculations indicate that bbp is dominated by submicron particles, but in field populations bbp likely has a significant tail in the phytoplankton size domain.
Satellite bbp covaries with POC
Loisel, H., E. Bosc, D. Stramski, K. Oubelkheir, and P.-Y. Deschamps (2001) Geophys. Res. Lett., 28:4203-4206.Stramski, D., R. A. Reynolds, M. Kahru, and B. G. Mitchell (1999)
Science, 285:239-242.
Therefore – bbp should be a better estimate of phytoplankton carbon than POC
If true – then the ratio of chlorophyll:bbp should track phytoplankton Chl:C
and thus phytoplankton growth rates ( )
{once a correction of the background bacterial population is accounted for}
(10)
(11)
(12)
(13)
(14)
0.1 1 10 100 1000 m
Practical POC
Particulate beam attenuation (cp)
Particulate backscatter (bbp)
-
0 5 10 15 20 25 300.01
0.04
0.07
0.10
0.13
0.16
0.19
0 1 2 3
0.005
0.020
0.035
0.050
0.065
0.080
0.20.40.60.81.00.001
0.006
0.011
0.016
Chl
:C (m
g m
g-1 )
Light (moles m-2 h-1)
Temperature (oC)
Chl
:Cm
ax
Growth rate (div. d-1)
Chl
:Cm
in
Low Nutrient stress High
3 primary factors
Light
Temperature
Nutrients
Chl:C physiologyLa
bora
tory
Chl:Cmax
Chl:Cmin
Dunaliella tertiolecta20 oCReplete nutrientsExponential growth phase
Geider (1987) New Phytol. 106: 1-34
16 species = Diatoms
= all other species
Laws & Bannister (1980) Limnol. Oceanogr. 25: 457-473
Thalassiosira fluviatilis = NO3 limited cultures
= NH4 limited cultures
= PO4 limited cultures
-
0.0 0.2 0.4 0.6 0.80.000
0.001
0.002
0.003
0.004
0.005
75o
0o
15o
30o
90o
60o 45o
60o
75o90o
15o
30o
45o
75o
0o
15o
30o
90o
60o 45o
60o
75o90o
15o
30o
45o NP
SP SA
NANP
SP
CP
SA
NA
SI
NI CA
SO
L0L1L2L3L4
SO-all
Chlorophyll
Variance L
evel
excluded
28 Regional Bins
b bp (
m-1
)
Chlorophyll (mg m-3)
‘physiology domain’
‘biomass domain’
Intercept = bbp from stable component of the bacterial population
C = scalar (bbp – intercept) = 13,000 (bbp – 0.00035)
Intercept = 0.00017 m-1 Stramski & Kiefer (1991) Prog. Oceanogr. 28, 343-383 Cho & Azam (1990) Mar. Ecol. Prog. Ser., 63, 253-259
Phytoplankton Carbon = 25 – 35% POC Eppley et al. (1992) J. Geophys. Res., 97, 655-661 DuRand et al. (2001) Deep-Sea Res. II, 48, 1983-2003 Gundersen et al. (2001) Deep-Sea Res. II 48, 1697-1718
-
0
0.1
0.2
0.3
0.4
0.5
0.6
0 0.2 0.4 0.6 0.8
0.0 0.2 0.4 0.6 0.80.000
0.001
0.002
0.003
0.004
0.005b b
p (m
-1)
Satellite bbp vs field cp
Same bilinear pattern
Anticipated intercept bbp > 0 cp = 0
Ratio of slopes = 0.0008
{phytoplankton = 0.0005 – 0.0009}c p (m
-1)
Chlorophyll (mg m-3)
South Atlantic
-
0 5 10 15 20 25 300.01
0.04
0.07
0.10
0.13
0.16
0.19
0 5 10 15 20 25 300.01
0.02
0.03
0.04
0.05
0 1 2 3
0.005
0.020
0.035
0.050
0.065
0.080
0 1 2 3
0.005
0.007
0.009
0.011
0.013
0 5 10 15 20 25 300.001
0.006
0.011
0.016
0.20.40.60.81.00.001
0.006
0.011
0.016
Chl
:C (m
g m
g-1 )
Light (moles photons m-2 h-1)
Temperature (oC)
Chl
:Cm
ax
Growth rate (div. d-1)
Chl
:Cm
in
Low Nutrient stress High
Labo
rato
ry
Temperature (oC)
Low Nutrient stress High
Chl:C
(mg m
g-1)
Chl:C
max
Chl:C
min
Space
{median r2 = 0.85}
-
75o
0o
15o
30o
60o 45o
60o
75o90o
15o
30o 45o
75o
0o
15o
30o
60o 45o
60o
75o90o
15o
30o 45o NP
SP SA
NANP
SP
CP
SA
NA
SI
NI CA
SO90o90o
0.02
0.03
0.04
0.05
0.06
0.003
0.005
0.007
0.08
0.10
0.12
0.14
0.007
0.010
0.013
0.016
0.04
0.07
0.10
0.13
0.005
0.008
0.011
0.014
0.0
0.1
0.2
0.3
0.4
0.01
0.02
0.03
1998 1999 2000 2001 2002
0.10
0.15
0.20
0.25
0.30
0.008
0.012
0.016
0.020
Year
Phyt
opla
nkto
n C
hlor
ophy
ll (
) a
nd sc
aled
Car
bon
(
)
Satellite Chl:C
Ratio ( )
The 5 basic patterns
-
Growth rate ( ) = max f (nuts, temp) g (light)
2 divisions d-1
Banse (1991)
Chl:Csat
( Chl:CN,T-max ) Light
range = 0 to 1 range = 0 to 1
0 1 2 30.00
0.01
0.02
0.03
0.04
0.05
Light (moles photons m-2 h-1)
Chl
:C (m
g m
g-1 )
1 – exp -3light
-
0 0.5 1.0 1.5 2.0
Growth rate (division d-1)
Boreal Summer
Boreal Winter
Growth rate (divisions d-1)0.0 0.5 1.0 1.5 2.0
Rel
ativ
e Fr
eque
ncy
0.0
0.2
0.4
0.6
0.8
1.0
Rel
ativ
e fr
eque
ncy
-
Bor
eal W
inte
r
Chlorophyll basedCarbon based
Bor
eal S
umm
er
Net Primary Production (mg m-2)0 1200800400 1500
-
Backup Slides
-
0 5 10 15 20 250
5
10
15
20
25
0
5
10
15
20
25
0 5 10 15 20 25
Balch et al. 1992 Antoine et al. 1996
Megard 1972 Behrenfeld & Falkowski 1997
Net primary production (NPP) = f [biomass, physiology, g (I0)]
NPP = f [chlorophyll, Zeu, Pbopt, g (I0)]
{0.02 to 30 mg m-3}
{120 m}
{0 to 30 mg C mgChl-1 h-1}
{0 to 1}
{2 to 300 mg m-2}
NPP = f [Carbon, Zeu, , g (I0)]
{5 to 100 mg m-3} {0 to 2 div. d-1}
Measured Pbopt
Mod
eled
Pb o
pt40% 40-50%
Model #1
Model #4Model #3
Model #2
-
Growth Irradiance0 1 2 3
Phot
osyn
thes
is
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
Ligh
t-sat
urat
ed P
hoto
synt
hesi
s (P
* max
)
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
0.01 0.03 0.05 0.070.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
Light-limited Photosynthesis (Nmax)0.01 0.03 0.05 0.07 0.01 0.03 0.05 0.07
Ek = 146r2 = 0.63
n = 17o.d. = 0.2 - 0.5
Ek = 138r2 = 0.73
n = 14o.d. = 0.7 - 1.0
Ek = 118r2 = 0.95
n = 16o.d. = 1.4 - 1.9
Ek = 95r2 = 0.91
n = 16o.d. = 2.1 - 2.6
Ek = 59r2 = 0.79
n = 20o.d. = 2.7 - 3.3
Ek = 42r2 = 0.80
n = 19o.d. = 3.6 - 4.3
Ek = 27r2 = 0.75n = 17o.d. = 4.5 - 5.0
Ek = 21r2 = 0.50n = 14o.d. = 5.3 - 5.8
Ek = 12r2 = 0.77n = 25o.d. = 6.2 - 10.6
Optical Depth0 1 2 3 4 5 6 7
E k
0
40
80
120
160
Observed
Calculated
CBA
D
IHG
FE
J K
-
Sequential hours of Experiment0 20 40 60 80
Car
bon
Fixa
tion
& Ac
cum
ulat
ion
Rat
e
-6
-4
-2
0
2
4
6
Time of Day0 4 8 12 16 20 24
FLS
0.10
0.12
0.14
0.16
0.18
0.20
N:C
0.15
0.17
0.19
0.21
Time of Day0 4 8 12 16 20
rbcL
0
5
10
15
Sequential hours of Experiment0 20 40 60 80
Pbm
ax
0
2
4
6
8
10
12
"b
0.00
0.01
0.02
0.03A
D
CB