Deep-Sea Research II 47 (2000) 1077}1106

A model of plankton dynamics for the coastal

upwelling system of Monterey Bay, California
Rafael A. Olivieri, Francisco P. Chavez*
Monterey Bay Aquarium Research Institute, P.O. Box 628, 7700 Sandholdt Road, Moss Landing,
CA 95039-0628, USA
Received 28 August 1998; received in revised form 12 April 1999; accepted 11 May 1999

Abstract
A seven-box plankton model for the mixed layer of the open ocean (Fasham et al., 1990,
Journal of Marine Research 48, 591}639) was modi"ed and applied to the coastal upwelling
ecosystem of Monterey Bay, California. Upwelling velocities, photosynthetically active radiation and changes in mixed-layer depth drove the model. The observed long-term mean
(1989}1995) seasonal cycles of nitrate concentration, primary production and phytoplankton
concentration within the mixed layer were reproduced. Bacteria and zooplankton biomass, and
ammonium concentration were within the same order of magnitude as observations. Modeled
dissolved organic nitrogen (labile) values were an order of magnitude lower than observations
(total). New production estimates were high, leading to an annual f-ratio of 0.84, typical of
productive coastal upwelling systems. Export production was much lower, and horizontal
export by upwelling-induced advection was the dominant loss term for phytoplankton primary
production. It exceeded sinking and grazing and the #ux of zooplankton-derived detritus
throughout most of the year. The model suggests that a large proportion of the daily
phytoplankton growth is ungrazed in the mixed layer, and is advected downstream and o!shore
of the production site. ( 2000 Elsevier Science Ltd. All rights reserved.

1. Introduction
Coastal upwelling regions represent a small percentage of the world oceans surface,
but account for a disproportionately large fraction of the oceans primary production
(PP) and "sh yields (Ryther, 1969; Walsh, 1976; Chavez and Toggweiler, 1995).
Primary production in coastal waters is typically 3}5 times higher than open ocean
* Corresponding author: Fax: 001-831-775-1620.
E-mail address: chfr@mbari.org (F.P. Chavez)
0967-0645/00/$ - see front matter ( 2000 Elsevier Science Ltd. All rights reserved.
PII: S 0 9 6 7 - 0 6 4 5 ( 9 9 ) 0 0 1 3 7 - X

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R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

waters on a per unit of area basis, but export production, organic carbon reaching the
benthos, and accumulation of organic carbon in the sediments can be many times
larger (Berger et al., 1989). While the coastal upwelling process and its biological and
chemical consequences have been studied extensively (see Barber and Smith, 1981a,b;
Hutchings et al., 1995), these studies have been sporadic in space and time. Input
parameters needed for modeling coastal upwelling ecosystems, therefore, have been
only marginally available, and long-term data sets for model validation have been
lacking. In 1989 the Monterey Bay Aquarium Research Institute (MBARI) began an
intensive study of the coastal upwelling system of central California (Chavez, 1996),
incorporating biweekly to monthly ship expeditions together with continuous observations from strategically placed moored platforms (Chavez et al., 1997). The results
from these time-series measurements have provided insights into the processes regulating biological production in a coastal upwelling ecosystem as well as data for model
input and validation.
In this contribution we describe the modi"cation and application of an upper
open-ocean ecosystem model to a coastal upwelling ecosystem. The model we selected
was the Fasham et al. (1990) model, here referred to as the `FDMa model (Sarmiento
et al., 1993; Haney and Jackson, 1996). The FDM model is one of a series of
Phytoplankton}Nutrient}Zooplankton (PNZ) models that trace their roots to the
early work by Fleming (1939) and Riley (1946). The original FDM model was used
to simulate the seasonal cycle of nutrients and primary production at Bermuda's
Station S. More recently it has been incorporated into the Princeton North Atlantic
General Circulation Model (Sarmiento et al., 1993), and in a tropical Paci"c
ocean circulation model (Toggweiler and Carson, 1995) with reasonable success.
FDM also has been used as a test-bed for parameter optimization protocols (Matear,
1995), and to explore the consequences of variations in phytoplankton growth
formulations (Haney and Jackson, 1996). We explore the use of FDM for modeling
a coastal upwelling pelagic ecosystem. The primary goal of our modeling exercise was
to simulate the long-term mean (1989}1995) seasonal cycle of primary production
and then to explore the model predictions for other variables, in particular new and
export production.

2. Environmental setting
Monterey Bay is a complex and dynamic coastal upwelling environment (Breaker
and Broenkow, 1994; Rosenfeld et al., 1994; Paduan and Rosenfeld, 1996), and
substantially di!erent from the oligotrophic subtropical open ocean around Bermuda
where the FDM model was originally applied. The ocean around Bermuda's Station
S, even though more variable than previously thought (Goldman, 1988), is closer to
steady state than coastal upwelling ecosystems (Mann and Lazier, 1991). Monterey
Bay #uctuates seasonally from high primary production (PP) during spring and
summer upwelling, to a relative low level during winter non-upwelling periods
(Chavez, 1996). The euphotic zone integrated PP for Monterey Bay during the winter
low (Chavez, 1996) is similar to the mean PP reported for the Hawaii Ocean

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1079

Time-Series (HOT) site (Karl and Lukas, 1996), and the Bermuda Atlantic TimeSeries (BATS) site (Siegel et al., 1995). During the spring and summer values reach
levels similar to those reported for other coastal upwelling environments (Chavez and
Barber, 1987). Skogsberg (1936) and Bolin and Abbott (1963) used physical and
biological characteristics to describe three major oceanographic seasons: upwelling
from about March through July, oceanic during August and September, and the
winter Davidson period (see Pennington and Chavez, 2000, for details).
2.1. The observations
The sampling program at MBARI has included stations throughout the Monterey
Bay region. We focused on the region near the M1 mooring (Fig. 1) because this

Fig. 1. Sea surface temperature image of the Monterey Bay region during the upwelling season of 1995.
Shown is the site of the mooring M1 located in the plume of recently upwelled water, where the modeling
e!ort was focused.

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R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

station has been sampled continuously since the beginning of the program in mid1989. It is also a location where a mooring (Chavez et al., 1997) collects continuous
measurements of key input variables for the model. This site is close to an area of
active upwelling. At the onset of upwelling the coolest waters are found north of the
bay (Rosenfeld et al., 1994). This colder upwelling center may be the source of the
water found downstream at the M1 site, although it is likely that upwelling occurs
along the entire mouth of the bay. During the upwelling period drifters deployed in
the upwelling center reach the M1 site in two days (Chavez et al., 1997). The
observations made on the biweekly to monthly ship time-series are described in
greater deal elsewhere (Chavez, 1996; Pennington and Chavez, 2000). The primary
observations used to verify the model results were primary production and nitrate
concentration.

3. The model
The FDM model is a seven-box plankton model of the oceanic mixed layer (Fig. 2,
Tables 1 and 2). The model has three living compartments de"ned as phytoplankton
(P), zooplankton (Z), and bacteria (B); and four nonliving ones: nitrate (N ),
/
ammonium (N ), dissolved organic nitrogen (DON) and detritus (D). The main
3
physical parameters are mixed-layer depth and photosynthetically active radiation
(PAR). The details of FDM can be found in the original publication (Fasham et al.,
1990). Below we describe only the modi"cations used in the present set of
simulations.

Fig. 2. Box model of the planktonic food web used to represent the Monterey Bay ecosystem. The direct
#ow from phytoplankton to detritus was eliminated from the original FDM model (see text for details).
Parameters were adjusted for an upwelling environment, and a new algorithm was used to calculate nitrate
input.

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Table 1
List of original major equations for the seven compartments of the FDM model
Phytoplankton
dP
(m#h`(t))P
"(1!c ) p(t, M, N , N )P!G !k P!
1
/ 3
1
1
dt
M

(1)

Zooplankton
dZ
h(t)Z
"b G #b G #b G !k Z!k Z!
1 1
2 2
3 3
2
5
dt
M

(2)

Bacteria
dB
(m#h`(t))B
"; #; !G !k B!
1
2
2
3
dt
M

(3)

Detritus
dD
(m#h`(t)#S )D
D
"(1!b )G #(1!b )G !b G !k D#k P!
1 1
2 2
3 3
4
1
dt
M

(4)

Nitrate
(m#h`(t))
dN
/ "!J(t, M)Q (N , N )P#
(N !N )
1 / 3
0
/
M
dt

(5)

Ammonium
(m#h`(t))
dN
3 "!J(t, M)Q (N )P!; #k B#(ek #(1!X)k Z!
N
2 3
2
3
2
5
r
M
dt

(6)

Dissolved organic nitrogen

dN
(m#h`(t))
$ "c J(t, M)Q (N , N )P#k D#(1!e)k Z!; !
N
1
2 / 3
4
2
1
$
dt
M

(7)

3.1. Upwelling velocities

FDM used a constant di!usion rate (m) of 0.1 m d~1 to supply the mixed layer with
nutrients. However, for Bermuda the primary source of nutrients was associated with
changes in mixed-layer depth. During winter, convection mixed subthermocline
waters into the surface layer. The original FDM model did not support upwelling. For
the coastal upwelling simulations an array of upwelling velocities was used to
transport dissolved material into the mixed layer. Daily mean upwelling from a 12-yr
data set of upwelling indices (Mason and Bakun, 1986) was computed for the
Monterey Bay region. Values ranged from !55 to 207 m3 s~1 100 m~1 of coastline.
These were then divided by an estimate of the Rossby radius of deformation of 32 km
at 363N (Chavez, 1996). The results were vertical velocities (m ) that #uctuated from
5
!1.5 (downwelling) to 5.6 m d~1 (upwelling) (Fig. 3A). These upwelling velocities fell
within the range of values of 1}10 m d~1 for coastal upwelling systems (Smith, 1968),

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R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

Table 2
List of variables from the seven major equations of FDM model
c
1
p
t
M
G
1
k
1
m
h(t)
h`(t)
b
1
b
2
b
3
G
2
G
3
k
2
k
5
;
1
;
2
k
3
S
D
k
4
J
Q
e
X

DON fraction exudated by phytoplankton

Average phytoplankton growth in a day
Time
Mixed-layer depth
Loss of phytoplankton by zooplankton grazing
Natural mortality rate of phytoplankton (senescence)
Vertical di!usive mixing rate
Change in mixed-layer depth through time, used for swimmers
Maximum value between h(t) and 0; used for non-swimmers
Zooplankton assimilation e$ciency of phytoplankton
Zooplankton assimilation e$ciency of bacteria
Zooplankton assimilation e$ciency of detritus
Loss of bacteria by zooplankton grazing
Loss of detritus by zooplankton grazing
Zooplankton speci"c excretion rate
Zooplankton speci"c mortality rate
DON uptake by bacteria
Ammonium uptake by bacteria
Bacteria speci"c excretion rate
Detritus sinking rate
Detritus speci"c breakdown rate
Light limited growth rate
Nutrient limitation parameter
Fraction of zooplankton excretion as ammonium
Fraction of zooplankton mortality to detritus

had the same dimensions, but were an order of magnitude greater than the vertical
di!usion rate (m) of the original FDM model.
The coastal upwelling system was divided conceptually into two layers. The upper
(surface) layer represents the shallow mixed layer with phytoplankton, primary
production and nutrients; the low (subsurface) layer represents the upwelling source
waters with no phytoplankton and high nutrients (Barber and Smith, 1981a,b). In our
model water that upwelled resulted in the removal of an equal volume from the mixed
layer, and therefore an export of unused nutrients and primary production. Downwelling was assumed to cause loss of material from the mixed layer. The model
neglects primary production that occurs below the mixed layer and input of phytoplankton. Data for Monterey Bay indicate that the majority of the total annual primary
production occurs within the shallow mixed layer (Pennington and Chavez, 2000).
3.2. Par values
FDM calculated photosynthetically active radiation (PAR) at noon from an oktas
array (a measure of cloudiness), ratio of PAR to solar irradiance, latitude, time of year,
and transmittance through the air}surface water interface (Fasham et al., 1990). Daily
PAR was available for Monterey Bay from LI-COR Inc. sensors on the M1 mooring
(Chavez et al., 1997) avoiding the need for calculation of PAR from oktas. Daily PAR

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1083

Fig. 3. (A) Mean daily upwelling velocities calculated from a 12 yr data set of daily upwelling indices for
363N 1223W (see text); (B) annual cycle (1992}1995) of photosynthetically available radiation (PAR)
collected at the M1 mooring; (C) annual cycle (1989}1995) of daily mixed-layer depth calculated from CTD
pro"les and using a of 0.13C; (D) annual cycle (1989}1995) of sea surface temperature (SST) at M1.

values at noon from 1992 to 1995 were used to calculate average noon values for every
Year Day (YD). These average daily noon values of PAR were smoothed with a 7-day
running mean (Fig. 3B) and used as the annual cycle of solar irradiation after
multiplying by 0.96 for air}water surface interface transmittance (same value used by
FDM).
3.3. Sinking rates
Constant sinking rates were assigned to phytoplankton (S ), and detritus (S ), but
P
D
were corrected at every time step with the upwelling velocities (m ) (Malone, 1971;
5
Barber and Smith, 1981a,b). This modi"cation changed the last term of Eq. (1)
(phytoplankton) and Eq. (4) (detritus). For example, for detritus it changed to
h`(t)D#E(t)D
!
,
M

(8)

where E(t) is the export resulting from the interaction of sinking and upwelling
velocities at any given moment de"ned as
E(t)"max(m , S )!min(m , S ).
5 D
5 D
The last term in the phytoplankton Eq. (1) is analogous to Eq. (8).

(9)

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R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

3.4. Mixed-layer depth

A * of 0.13C with respect to SST was used to calculate the mixed-layer depth
(MLD) (Dickey, 1991); in the literature * ranges from 0.02 to 0.53C (Husby and
Nelson, 1982; Marra et al., 1990; Lentz, 1992). CTD data collected between 1989 and
1995 was used to build an average annual cycle of MLD for Monterey Bay (Fig. 3C).
3.5. Sea-surface temperature (SST )
SST measured daily with a CTD on the M1 mooring, and on the biweekly CTD
cruises, was used to calculate average daily SST for every YD. As with PAR, the
average daily SST values were smoothed with a 7-day running mean. SST was used to
calculate nitrate below the mixed layer (N ), and to set the maximum phytoplankton
0
growth rate (see below) (Fig. 3D).
3.6. Nitrate concentration below the mixed layer (N )
0
In order to calculate N , the temperature of the water below the mixed layer was
0
converted to nitrate using a nitrate}temperature relationship (Fig. 4). The temperature below the mixed layer was calculated from thermocline strength and SST.
Thermocline strength is the di!erence between SST and temperature at the base of the
thermocline, and for Monterey Bay ranges from around 13 (winter) to 23C (summer)
(Husby and Nelson, 1982). For the simulations we used a thermocline strength that

Fig. 4. Second-order regression of nitrate versus temperature. Samples collected at stations H3 and M1
mooring from 1989 to 1995 and from depths between 10 and 60 m.

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1085

gradually increased from 1.253 (late winter) to 23C (late summer), re#ecting winter
vertical mixing and summer strati"cation. For any YD, the thermocline strength was
subtracted from SST, and this temperature was used in the nitrate}temperature
regression to approximate N . At temperatures *14.853C, N was set to
0
0
0.0 mmol m~3.
3.7. Phytoplankton terms
In FDM, < , the maximum phytoplankton growth rate was a constant, and was the
P
mean of the phytoplankton maximum growth calculated using the temperature
function of Eppley (1972). We calculated < every YD from SST using Eppley's (1972)
P
function. FDM used a value for a of 0.025 d~1 (W m~2)~1. Kudela (1995) reported
a values for Monterey Bay that ranged from 0.015 to 0.031 mg C (mg Chl
a)~1 h~1(lE m~2 S~1)~1. With a 1 : 1 ratio of mol N to g Chl (Eppley et al., 1992) and
a Red"eld C : N ratio, these values ranged from 0.021 to 0.043 (W m~2)~1 d~1.
Measurements (F. Chavez, unpublished) during spring of 1995 ranged from 0.015 to
0.146 (W m~2)~1 d~1, with a mean a of 0.055 (W m~2)~1 d~1. Harrison et al. (1981)
values for the Peru upwelling system ranged from 0.003 to 0.099 (W m~2)~1 d~1, with
a mean of 0.042 (W m~2)~1 d~1. Harrison et al. (1981) also noticed that a increased
with Chl and proximity to coast. In our model we used an a of 0.03 d~1 (W m~2)~1,
slightly higher than the one used in FDM.
The phytoplankton growth equation used by FDM requires values for the attenuation coe$cients for phytoplankton and water (plus detritus and dissolved colored
material) k , and k (Fasham et al., 1990). k and k can be approximated with Riley's
#
8
#
8
(1946) linear equation:
k"k #k C,
(10)
8
#
where the intercept k is the light attenuation coe$cient of water plus the non8
phytoplankton and dead phytoplankton components, k is the attenuation by live
#
phytoplankton, C is chlorophyll concentration, and k is total extinction coe$cient
(Bannister, 1974). From chlorophyll and attenuation coe$cients (Fig. 5) we calculated
a k of 0.146 m~1 and k of 0.024 m2 (mg Chl a)~1. For a concentration of 0.5 mg
8
#
Chl m~3, typical of low chlorophyll waters, Eq. (10) estimates a k of 0.158 m~1, almost
identical to the k of 0.15 m~1 suggested for clear coastal water by Clarke and Denton
(1962). The value estimated for k is similar to that reported by others (Bannister,
#
1974; Fasham et al., 1990).
Dortch (1990) showed that ammonium inhibition was variable, and proposed that
a combination of preference and ammonium inhibition better described the ammonium}nitrate interaction. In Monterey Bay, Berges et al. (1995) and Kudela (1995)
did not "nd evidence for ammonium inhibition at ambient ammonium concentrations. Thus, ammonium inhibition was eliminated as recommended by Haney and
Jackson (1996) and the nitrogen growth function changed to
N
N
/
/
Q(N N )"Q (N )#Q (N )"
#
/, 3
1 /,
2 3
K 3#N
K 4#N
NO
NH
/
3

(11)

1086

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

Fig. 5. Linear regression of extinction coe$cient (k) calculated from secchi disk versus mean chlorophyll-a
concentration (C) in the euphotic zone. The slope of the line is the attenuation by chlorophyll-a (k ). The
#
intercept is the attenuation by seawater and dissolved and suspended material (k ).
8

FDM used a half-saturation constant of 0.5 mmol m~3 for K 3 and K 4 . A few
NO
NH
K 3 values were available for Monterey Bay, that ranged from under 1 to
NO
4 mmol m~3 (Kudela, 1995). To incorporate ammonium preference over nitrate
(Kudela, 1995), we set K 3 at 0.75 mmol m~3, and K 4 at 0.5 mmol m~3, because
NO
NH
of the generally low ammonium concentrations measured in Monterey Bay (Berges
et al., 1995 C. Sakamoto pers. comm.).
FDM used a natural mortality factor (k ) not included in the grazing mortality,
1
which represented a direct #ow of dead phytoplankton cells to the detritus pool within
the mixed layer. The use of k has been questioned (Haney and Jackson, 1996) and we
1
removed it. We used a phytoplankton sinking rate (S ) of 0.5 m d~1 (see sinking
P
section above), a value between the 1 m d~1 for coastal diatoms growing under
nutrient replete conditions, and the 0.2 and 0.4 m d~1 for dino#agellates and small
diatoms (Bienfang and Harrison, 1984).
3.8. Zooplankton terms
We changed three aspects of the FDM zooplankton equation. The "rst two are part
of the grazing rate function (G), and the third one modi"ed the zooplankton mortality
function. FDM de"ned the grazing of phytoplankton (or any prey) by
gZp P2
1
G "
,
1 K (p P#p B#p D)#p P2#p2B2#p D
Z 1
2
3
1
3

(12)

where (g) is the zooplankton maximum speci"c grazing rate, (K ) is the half-saturation
Z
constant for zooplankton grazing, and p is the preference as function of relative prey
1
density. FDM used 1 d~1 for g, adequate for zooplankton communities of oligotrophic waters dominated by small copepods and heterotrophic protists. For coastal
waters with larger predators, g should be lower. For example, a maximum speci"c

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1087

ingestion rate of around 0.2 d~1 was calculated for Euphausia lucens (Stuart, 1986), the
dominant euphausiid and contributor of about half of the zooplankton biomass in the
Benguela current (Stuart, 1986). Euphausiids are also abundant in Monterey Bay;
therefore g was set at 0.6 d~1. A plot of carbon-speci"c grazing rates and diatom
carbon concentration (Stuart, 1986) showed the half saturation grazing rate for
E. lucens feeding on diatoms to be around 220 lg C l~1, which by Red"eld conversion
equates to 2.76 mmol N m~3. We, therefore, increased K to 3 mmol N m~3 from the
Z
1 mmol N m~3 used in FDM.
The "nal modi"cation to the zooplankton equation was to the mortality induced by
predators outside the model (k Z), used as a closure term in the equation. With the
5
original function primary production goes into an unrealistic cyclic behavior under
nutrient rich conditions (Steele and Henderson, 1992). The mortality function was
changed to a quadratic (k Z2) as recommended by Steele and Henderson (1981). Fasham
5
(1995) and McGillicuddy et al. (1995) utilized quadratic functions for the same purpose.
3.9. Bacteria, detritus and dissolved organic nitrogen terms
With the exception of an increase in S (see sinking topic above) from 10 to
D
25 m d~1, we kept the inputs for B, D, and DON as in FDM. The change of S was
D
based on a 25 m d~1 sinking rate for small fecal pellets calculated by Smith (1987), and
the 20}30 m d~1 rates measured in situ in Monterey Bay for small marine snow
aggregates (Pilskaln et al., 1996).
3.10. Model validation
Primary production (PP) integrated over the euphotic zone and nitrate concentration in the mixed layer from Monterey Bay were used to validate the model results. An
average annual curve of daily PP was built from PP measured from 1989 to early
1996. A similar curve was constructed for nitrate concentration in the mixed layer
from values obtained from 1989 to 1995. We used Red"eld ratios to convert model
PP, in units of mmol N m~2 d~1, to mg C m~2 d~1, for comparison with the 14C incubations. We followed the recommendation of Fasham et al. (1990) and (1) used
a time step of 0.2 days, and (2) ran the model for various years (at least 4) to ensure
a stable annual pattern.
4. Results
4.1. Evolution of the MB model
The "rst simulation was as speci"ed by the original FDM model with the PAR
array, instead of oktas. Values estimated for Monterey Bay were used for
k , k , a, K 3 , and MLD. < was set at 1.9 d~1 from average SST. N was set at
8 #
NO
P
0
a constant 10 mmol m~3. PP oscillated in a cyclic manner, as predicted by Steele and
Henderson (1992), and did not resemble the "eld measurements (Fig. 6A). N also did
/

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Fig. 6. Annual cycle of integrated primary production (A, C, E, G, I, K) and corresponding nitrate
concentration (B, D, F, H, J, L) for a series of simulations, starting with the original parameterization of the
FDM model conditions to the "nal con"guration for the MB model, see text for details.

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1089

not resemble the "eld data (Fig. 6B). An array of upwelling velocities (m ) was included
5
instead of vertical di!usion (m), a thermocline strength of 1.53C was used to calculate
N , S was corrected at every time step with the corresponding m , k and t were
0 D
5 1
removed, and g was lowered to 0.6 d~1 for the next simulation. PP maintained a cyclic
behavior, but the peaks reached higher values of around 1,000 mgC m~2 d~1
(Fig. 6C). This simulation produced N higher than the observed "eld measurements
/
but with a similar seasonal cycle (Figs. 6D and 7B).
The quadratic zooplankton mortality function was incorporated and the highfrequency cyclic behavior in PP was eliminated (Fig. 6E). A value of 0.05 mmol~1 d~1
was used for k ; it removed the high-frequency oscillations but did not fully suppress
5
zooplankton dynamics. In this simulation PP increased during the upwelling season
to more than 4000 mgC m~2 d~1, almost twice as high as the observed mean (Chavez,
1996). In contrast, PP during late summer was lower than the "eld measurements. The
high PP resulted in a marked decline in N , (Fig. 6F).
/

Fig. 7. (A) Integrated primary production integrated (mixed layer) simulated by the MB model and average
(1989}1996) integrated primary production (euphotic zone). (B) As in (A) but for nitrate concentration in the
mixed layer.

1090

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

In the next simulation we increased K to 3 mmol m~3. This modi"cation reduced

Z
the slope of the Michaelis}Menten grazing curve, resulting in a decrease of zooplankton grazing pressure, and preventing the collapse of PP at the end of the
upwelling season (Fig. 6G). However, PP was still twice that observed in the "eld
resulting in lower than observed N (Fig. 7G).
/
Addition of sinking rate (S ) of 0.5 m d~1, increasing S to 25 m d~1, and using daily
P
D
SST to calculate < had minimal e!ect on PP or N (not show). A variable thermocline
P
/
strength resulted in a slight increase in PP between YD 122 and 305, with the largest
change between about YD 200 and 250 (Fig. 6I). The increases in PP occurred without
a further reduction of N (Fig. 6J). For the last simulation we decreased a to 0.03 d~1
/
(W m~2)~1. This last simulation (Table 3), designated the `MB modela (Monterey
Bay), gave what we considered the most realistic simulations of PP and N , (Fig.
/
6 K and L, Figs. 6K and L, 7 A and B) and provided the basis for exploring the #ows
between boxes as well as the resulting biomass of the compartments.
4.2. Phytoplankton, primary production, export and f-ratios
The annual integrated PP estimated using the MB model was similar to the
observed multi-year PP mean (Table 4). The model annual cycle had higher values at
Table 3
Input values for MB model (Fig. 6K and L). (*) Same values as in FDM
c
1
S
P
m
I
0
N
0
b
1
b
2
b
3
k
2
k
5
g
<
P
a
K
8
K
#
K 3
NO
K 4
NH
K
Z
K
B
k
3
<
B
S
D
k
4
g
e
X

0.05
0.5 m d~1
Upwelling array (m d~1)
PAR array
Calculated from temperature array
0.75
0.75
0.75
0.1 d~1
0.05 (mmol N)~1 m3 d~1
0.6 d~1
Calculated from temperature array
0.03 (W m~2)~1 d~1
0.146 m~1
0.024 m2 (mmol N)~1
0.75 mmol N m~3
0.5 mmol N m~3
3.0 d~1
0.5 mmol N m~3
0.05 d~1
2.0 d~1
25 m d~1
0.05 d~1
0.6
0.75
0.33

*
*
A
*

*
*
*
*
*
*
*
*

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1091

Table 4
Annual averages for measured and simulated values from Monterey Bay and other coastal systems
Name

Units

Model

Phytoplankton
Zooplankton
Bacteria
DON
Ammonium
Nitrate
Detritus
Primary production
Export of primary production
Phytoplankton nitrate uptake
Phytoplankton ammonium uptake
Zooplankton grazing phytoplankton
Export of detritus
Productivity index

mmol N m~3
mmol N m~3
mmol N m~3
mmol N m~3
mmol N m~3
mmol N m~3
mmol N m~3
g C m~2 yr~1
g C m~2 yr~1
mmol NO m~3 d~1
3
mmol NH m~3 d~1
4
mmol N m~3 d~1
mol N m~2 yr~1
mg C mg Chl~1 d~1

3.78
1.54
0.60
0.07
0.14
3.24
0.06
472
287
1.48
0.23
0.50
0.51
31

Field values
2.61!
0.56"
1.03#
3.9$
0.49%
4.10%
459%
370&
0.79'
0.23)
0.48+
1.08&
28%

0.11+
0.18&

!Mean value from Monterey Bay stations H3 and Mooring M1 and mmol N : mg Chl a ratio of 1 (Eppley
et al., 1992).
"Acoustic estimated mean biomass for Monterey Bay (Chavez, unpublished).
#Mean biomass from #ow cytometry (Buck et al., 1996), 20 fg C per bacterium (Fuhrman, 1981), and 3 : 1
C : N ratio (Antia et al., 1991).
$One single value for 10 m reported for Monterey Bay by Hansell (1993).
%Mean value from Monterey Bay stations H3 and Mooring M1.
&From sediment traps in Monterey Bay; A) from euphotic zone, B) at 450 m (Pilskaln et al., 1996).
'15N uptakes and Model (Kudela and Chavez, 2000).
)15N uptakes from Monterey Bay (Kudela and Dugdale, 2000).
+From zooplankton mean biomass estimated with acoustics (Chavez unpublished) and a maximum
consumption/biomass ratio of (A) 0.86 d~1 for small copepods calculated from values for Peru upwelling
system (Dagg et al., 1980), and (B) 0.2 d~1 for euphausiids of the Benguela current (Stuart, 1986).

the start of the upwelling season and lower ones for the late summer months (Fig. 7A).
Winter values were around 500 mgC m~2 d~1 and increased quickly between YD 61
and 92, with the intensi"cation of upwelling and reduction of MLD, and then
decreased with the reduction in upwelling.
In the model, the water upwelled contained no phytoplankton. Thus, for each
parcel of water upwelled into the mixed layer an equal volume of water with
phytoplankton was removed. This process is analogous to net horizontal advection,
and was used to estimate the amount of PP exported horizontally. Upwelling drove
an o!shore export of 3.61 mol N m~2 yr~1 or 287 gC m~2 yr~1 of PP. During upwelling the o!shore export was between about 10 and 35 mmol N m~2 d~1 or 800
and 2780 mgC m~2 d~1, and was much larger than the export of detritus from the
system (Fig. 8A). During non-upwelling conditions, the export of PP and detritus were
similar.
The MB model estimated a phytoplankton concentration (P) of between 1 and
2 mmol N m~3 for the start of the year (Fig. 9A). P increased quickly to around
7}9 mmol N m~3 early in the upwelling season, just after YD 92, and remained

1092

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

Fig. 8. (A) Total export of phytoplankton and detritus from the mixed layer. (B) Plot of detritus export ratio
(e-ratio) versus primary production.

relatively high for about 50 days. P began to decrease until it reached its lowest values
of under 2 mmol m~3 by the next winter. Converting the N to chlorophyll using a 1 : 1
ratio of mol N to g Chl (Eppley et al., 1992), the modeled P was slightly higher, but
with a similar seasonal pattern as the "eld observations.
Phytoplankton uptake of N and N allowed the partitioning of PP into `newa and
/
3
`regenerateda production, respectively (Dugdale and Goering, 1967) and calculation
of the `f-ratioa, the ratio of new to total production. (Eppley and Peterson, 1979).
N uptake in the model was relatively low and constant throughout the year, but the
3
N uptake increased as a function of upwelling, and resulted in a f-ratio that changed
/
throughout the year (Fig. 10A). A plot of f-ratio versus PP produced a hyperbolic
distribution that increased quickly from 500 to 1000 mg C m~2 d~1, and plateau at
f-ratios between 0.8 to 0.9 after 1000 mg C m~2 d~1 (Fig. 10B). The model f-ratios
around 0.9 during high PP compared well to the measured f-ratio of 0.89 and PP of
1964 mgC m~2d~1 during the upwelling season of 1991 in Monterey Bay (Chavez
and Smith, 1995). We calculated a geometric mean (GM) regression of f-ratio versus

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1093

Fig. 9. (A) Annual cycle of biomass calculated by the MB model for bacteria, phytoplankton and
zooplankton plotted with mean (1989}1995) phytoplankton biomass (calculated from chlorophyll assuming a 1 : 1 ratio of mol N to g Chl (Eppley et al., 1992)). (B) NH and DON uptake by bacteria estimated by
4
the MB model.

the ratio of nitrate to nitrate plus ammonium from the model. The GM regression line
from the MB model was similar to that calculated by Harrison et al. (1987) for eight
coastal systems and the coast of Peru (Fig. 10C).
4.3. Zooplankton biomass and grazing
The model estimated zooplankton biomass (Z) to be around 1 mmol N m~3 in the
winter, increasing slightly along with P until it stabilized around 2 mmol N m~3

1094

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

Fig. 10. (A) Annual cycle of f-ratio from the MB model. (B) Hyperbolic distribution of f-ratio versus
integrated primary production for the same simulation. (C) Geometric mean (GM) regression of f-ratio
versus ratio of nitrate to total inorganic nitrogen (nitrate plus ammonium) concentration in mixed layer
calculated from MB model. For comparison, GM regression lines for data from eight coastal sites
(y"0.97x!0.09) and the coast of Peru (y"1.24x!0.33) (Harrison et al., 1987) are included.

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1095

(Fig. 9A). Silver and Davoll (1975,1976,1977) collected zooplankton samples

with a 330 lm net at a station located a few kilometers north of M1. They
reported their values in displacement volumes (ml 1000 m~3). We converted
these to mmol N m~3 assuming 1 ml of sample equal to 1 g wet-weight (Ahlstrom
and Thrailkill, 1963), 0.052 gC per g wet weight (Frost, 1987) and C : N mole ratio of
6.3 (Parsons et al., 1984). The lowest estimates of Z were about twice as the high as
those of Silver and Davoll (1975,1976,1977). The annual mean Z was higher but
similar in magnitude to the values estimated from acoustic data in Monterey Bay
(Table 4).
Zooplankton in the model grazed a phytoplankton biomass of 2.0 mol N m~2 yr~1
or 162 gC m~2 yr~1. Bacterial biomass grazed by zooplankton was (0.06 mol
N m~2 yr~1, or (3% of the phytoplankton biomass. The low detritus concentration
(about an order of magnitude lower than that of bacteria) resulted in a negligible
ingestion of detritus (fecal pellets) by zooplankton.
4.4. Bacteria
Bacterial counts were available from #ow cytometry samples from Monterey Bay
and were converted to biomass in mmol N m~3, using 20 fgC per bacterium (Fuhrman et al., 1989) and a C : N mol ratio of 3 : 1 (Antia et al., 1991). The bacterial
biomass (B) from the model was slightly lower than the biomass estimated by #ow
cytometry (Table 4). The standing stock of bacteria did not #uctuate greatly over the
course of the year (Fig. 9A). Highest bacterial integrated biomass was observed during
winter months, and lowest was from the end of upwelling and start of the oceanic
period. These #uctuations in standing stock numbers were presumably a consequence
of the change in MLD throughout the year.
The annual pattern in bacterial DON uptake (Fig. 9B) was similar to the annual
pattern of P. During times of high PP phytoplankton was the major contributor of
DON (Fig. 11A). In contrast, during times of low production, particularly the "rst
2 months and the last month of the year, zooplankton contributed more to the DON
fraction. The bacterial ammonium uptake followed the DON uptake (Fig. 9B), as in
the model ammonium is used by bacteria as a source of nitrogen (Fasham et al., 1990).
Ammonium was also produced by bacteria, but the contribution was minor when
compared with that of zooplankton.
4.5. Nitrate and ammonium
In the model the two forms of nitrogen available to phytoplankton were nitrate
(N ) and ammonium (N ). The model produced cycles that were out of phase
/
3
for N and N : summer had high N and low N , and winter had low N and high
/
3
/
3
/
N (Fig. 7B and 11B). The N calculated by the model was in good agreement with the
3
/
expected values, particularly during the upwelling season (Fig. 7B) (Table 4). Highest
N values were just under 15 mmol m~3 and occurred after the peak in upwelling
/
velocities. Lowest N values were from the end of the Davidson and the oceanic
/
season. The concentration of N in the mixed-layer resulted from a balance between
/

1096

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

Fig. 11. (A) Production of DON by zooplankton, phytoplankton and detritus estimated by the MB model.
(B) Annual pattern of ammonium, DON and detritus concentration estimated by the MB model.

upwelling, changes in the mixed-layer depth, and phytoplankton uptake. On a yearly

basis upwelling accounted for about 90% of the nitrate supply.
The model estimates the input of nitrate into the mixed layer using the di!erence
between N and N (Table 1, Eq. (5)). Therefore, the reported nitrate input into the
0
/
mixed layer is a net input, and the export by o!shore #ow is not directly calculated.

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1097

However, the o!shore nitrate #ow can be calculated as N times the upwelling
/
velocity. The calculated annual o!shore export of nitrate was 3.63 mol N m~2 yr~1
and 41% of the annual nitrate supply.
The supply of nitrate resulting from changes in mixed-layer depth was limited to
winter when the mixed-layer depth increases (Fig. 3C). On an annual basis this
contribution was less than 10%. However, the increase in mixed-layer depth is during
periods of weaker upwelling; therefore when it occurs, it is the dominant source of new
nitrogen.
N concentration calculated by the model was low during most of the year, and
3
reached its highest values of 0.45 mmol m~3 during winter (Davidson period)
(Fig. 11B). Comparison between observed and model values (Table 4) suggest that the
model underestimates this property. Low N , at or below 0.1 mmol m~3 for most of
3
the year, had the e!ect of controlling Q, the phytoplankton nitrogen-dependent
growth function. According to Eq. (11), under optimal nutrient conditions the maximum value of Q, approaches 2. In the model the highest Q values were under 1.
4.6. Detritus and DON
The detritus (D) and DON fractions in the model are byproducts from other
compartments. D originated from the non-assimilated fraction (b) of the zooplankton
grazing (which equates to fecal pellet production). DON originates from zooplankton
excretion that is not ammonium (1!e), the breakdown of detritus (k ), and the
4
exudation fraction (c) of phytoplankton production (Fasham et al., 1990).
The annual pattern in D concentration shows highest values during upwelling, but
the changes are extremely small and parallel to those of zooplankton (Fig. 11B).
A larger di!erence was found when the mixed-layer depth integrated D values were
compared. The highest values were associated with winter (Davidson), while the
lowest values occurred at the end of upwelling during the summer. There was,
therefore, no correlation between PP and detritus #ux.
On an annual basis the model predicts a detritus #ux out of the mixed layer of
about 9% of PP. Pilskaln et al. (1996) reported values of 18% PP #ux for the base of
the euphotic zone, and 3% measured #ux at 450 m. The detritus sinking rate was
always greater than the upwelling velocity, so detritus was not exported horizontally
but sank locally. In the model the e-ratio, or ratio of detritus export to total
production approximates a negative power function, with higher relative export
during low PP, and low relative ratios at higher PP (Fig. 8B). The MB model
predicted DON concentrations at least an order of magnitude lower than expected
(Hansell, 1993). The annual DON cycle was similar to that reported for other coastal
systems where the maximum values for dissolved organic matter tend to occur during
late summer (Williams, 1995). The MB model predicted the DON peak later in the
year than the peak of D (Fig. 11B). Highest DON concentrations were associated with
the cessation of upwelling, and they were about 10 times as high as those from winter.
The highest DON concentrations were associated with times of shallow MLD.
Highest DON values integrated over the mixed layer were about 4 times greater than
the lowest ones.

1098

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

5. Discussion
The FDM model was modi"ed and used to simulate the annual cycle of integrated
PP and nitrate concentration for the mixed layer of Monterey Bay (Fig. 7A). It also
produced results for phytoplankton, zooplankton and bacterial biomass that were
similar to the observations. Ammonium concentration was apparently underestimated, and DON was about an order of magnitude lower than the few observations. Several other rate parameters were similar to those measured in Monterey Bay
or reported in the literature (Table 4).
Construction of the MB model required few changes from the original equations
used in FDM suggesting that FDM may be a good representation of a generic pelagic
ecosystem. Changes were required in the basic physics (and therefore nutrient supply)
to account for the upwelling process. Model parameters were adjusted to account for
this process (i.e., phytoplankton sinking) and for the resulting food web structure
(diatom-dominated versus microbial loop). The use of a quadratic zooplankton
mortality function (Steele and Henderson, 1981) was important. It removed cyclic
oscillations in phytoplankton biomass and production under nutrient rich conditions
(Steele and Henderson, 1992) by decoupling phytoplankton and zooplankton growth
(i.e., diatom}macrozooplankton versus microbial loop). The quadratic mortality is in
essence the consumption of zooplankton by higher predators. It becomes important
at high zooplankton concentrations and places a cap on these populations.
The FDM model has now been applied with some success to open-ocean oligotrophic environments (Fasham et al., 1990), the mesotrophic equatorial Paci"c
(Toggweiler and Carson, 1995), the open-ocean North Atlantic (Sarmiento et al., 1993),
with some slight modi"cations to the subarctic Paci"c and Atlantic oceans (Fasham,
1995), and in this contribution to an eutrophic coastal upwelling system. The implication is that one core pelagic ecosystem model can be used for global simulations.
Previous e!orts dealt primarily with ecosystems that were microbial-loop-dominated;
our contribution has shown that it can also be applied to a coastal diatom-dominated
system. The challenge for global modelers will be to "nd (in their models) the physical
(or chemical) processes that lead to these two very di!erent pelagic ecosystems
(Chavez and Smith, 1995).
Current ecosystem modeling e!orts are incorporating additional phytoplankton
and zooplankton compartments in an e!ort to simulate the occurrence of these
di!erent ecosystems. The next level of complexity has two size classes of phytoplankton, one composed of small single-celled phytoplankton like Synechococcus, and the
other of larger phytoplankton like diatoms. Two size classes of zooplankton, one
composed of microzooplankton-like ciliates and the other of meso-and macrozooplankton, like copepods and euphausiids, are also required. The microzooplankton
graze primarily on the small single-celled phytoplankton and bacteria, while
the mesozooplankton primarily consume netphytoplankton, microzooplankton and
detritus. A drawback to increasing the number of boxes is that the connections or
#ows increase nonlinearly and therefore the model becomes more complex and
di$cult to balance. If it is primarily an either/or scenario (diatom}mesozooplankton
versus picoplankton}microzooplankton), then regulating the grazing term, in space or

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1099

time and as a function of micro- or macro-nutrient limitation, accomplishes the same

basic results without the computational overhead. Open-ocean iron fertilization
experiments (Coale et al., 1996; Kudela and Chavez, 1996) have highlighted the role of
iron in regulating ecosystem structure. The role of iron in regulating the ecosystem
structure of coastal upwelling systems only recently has been addressed (Chavez et al.,
1991; Hutchins and Bruland, 1998; Johnson et al., in press). Iron may play a controlling role in ecosystem structure on a global scale and be the critical switching
component, but further work is clearly required.
5.1. Primary production, new and export production
The model PP was very similar to the "eld observations (Fig. 7A; Table 4).
Measured and simulated values of the productivity index (primary production normalized to chlorophyll) display similar seasonal cycles (not shown) and similar
magnitudes (Table 4), lending further credibility to the PP simulation. Photosynthesis-irradiance and half-saturation nutrient uptake constants were parameters for
which additional "eld measurements are required. a and K 3 are most likely
NO
a function of nutrient concentration and species composition/cell size. K 3 is
NO
thought to increase with cell size (Malone, 1980), and within a species, clones from
coastal nutrient-rich waters tend to have higher K 3 than those from open ocean and
NO
nutrient-poor waters (Carpenter and Guillard, 1971). In Monterey Bay, the occurrence of larger phytoplankton cells, like chain-forming diatoms, is closely coupled to
the upwelling period (Fig. 12), when an increase of NO supply into the mixed layer
3
occurs (Bolin and Abbott, 1963; Malone, 1971; Garrison, 1976,1979; Schrader, 1981;
Chavez, 1996). The increase in nitrate favors the increase in the abundance of diatoms
during upwelling. Later, as nitrate becomes depleted, smaller cells with lower
K 3 would be favored.
NO

Fig. 12. Mean (1989}1995) spherical diameter of phytoplankton for samples collected at site M1 in
Monterey Bay.

1100

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

The model calculated an annual f-ratio of 0.84, with a new production of

5 mol N m~2 yr~1 or 400 g C m~2 yr~1. The model f-ratios of 0.9 during upwelling
high PP are supported by "eld values from Monterey Bay (Chavez and Smith,
1995; White and Dugdale, 1997). f-ratios higher than 0.5 are typical of coastal
upwelling systems dominated by diatoms (Hutchings et al., 1995). The model computed the f-ratio as the uptake of nitrate versus the uptake of nitrate and ammonium,
comparable to f-ratios calculated with 15N uptake experiments. Nitrate uptake is
a measure of new production (Dugdale and Goering, 1967), and therefore of the
carbon that should be exported vertically under steady state (Eppley and Peterson,
1979). Our estimates of detritus export (fecal pellets and comparable to what a sediment trap might catch) and the e-ratio (detritus export divided by PP) are signi"cantly
di!erent than new production and the f-ratio (Figs. 8B and 10B). The e-ratio versus
PP scatter plot (Fig. 8B) is similar to that measured by moored (Pilskaln et al., 1996)
and #oating sediment traps (J. Newton, personal communication). The model results
presented here are consistent with the observations, both of which show the lack of
a strong, positive correlation between PP and export in this highly productive
environment.
How do we explain the apparent paradox of new and export production? In the MB
model, a large fraction of the PP was unused within the mixed layer and exported
horizontally as ungrazed phytoplankton cells (Fig. 8A). The upwelling circulation
drove this horizontal export (which accounted for most if not all of the ungrazed PP
export). Similar mechanisms were invoked by Chavez (1996) to explain observations
of PP and nutrients during the 1992 El Nin8 o and by Pilskaln et al. (1996) to
explain the e-ratio trend determined from POC #ux and PP measurements. The
concept of a one-dimensional biological pump is clearly not appropriate for the
MB coastal upwelling system (Silver et al., 1998; Service et al., 1998) where there
is a space and time uncoupling between sources (nutrients, new production) and
sinks (vertical export). Recent modeling and experimental research in the openocean upwelling system of the equatorial Paci"c also show that horizontal
advection of new production needs to be considered to balance the production
budget (Landry et al., 1997; Loukos et al., 1998). The model results show that
nitrate uptake and carbon export are only equivalent when considered over appropriate temporal and spatial scales (Eppley, 1989; Legendre and Gosselin, 1989).
Further work is required to de"ne these scales for the coastal upwelling system
of MB.
5.2. Nitrate and ammonium
The model simulated the observed nitrate concentration (N ) accurately. N is
/
/
a function of physical supply and phytoplankton uptake. Physical supply is a function
of upwelling velocities, SST, thermocline strength, and change in mixed-layer depth.
SST and thermocline strength were used to estimate nitrate concentration (N ) below
0
the mixed layer. For future simulations N should come from (moored) measurements
0
of temperature or nitrate from below the mixed layer. The calculated horizontal
nitrate export was 40% of the physical supply, highlighting the importance of

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

1101

upwelling as a source of nutrients (and production) for o!shore communities (Chavez,

1996).
The climatological ammonium data set is small, and we know little about the
regeneration terms. The zooplankton component, which is the primary source of
ammonium, is also one of the least understood. We have not considered the supply of
ammonium to the mixed layer by upwelling or mixing. Grazing and excretion occur in
the upper waters of the mesopelagic zone, both in and below the mixed layer.
5.3. DON
The accurate measurement of DON in the ocean has been surrounded by controversy (Hansell et al., 1993; Bronk et al., 1994; Williams, 1995). Based on the values of
Hansell (1993) and Kudela (pers. comm.) the model underestimated surface DON
concentrations by an order of magnitude. However, this might be associated with
di!erences between measured (refractory) and modeled (labile) pools of DON. In the
model the total annual DON production by phytoplankton was much larger than the
DON produced by zooplankton (Fig. 11A). However, at times of low primary
production the zooplankton DON contribution was larger than the phytoplankton
contribution.
5.4. Zooplankton
One of the few multi-year zooplankton programs for Monterey Bay was that by
Silver and Davoll (1975,1976,1977). This program collected samples with a 330 lm
net. This net misses a signi"cant fraction of the zooplankton community (Hopcroft
et al., 1999). In the model, phytoplankton was the zooplankton main food source,
bacteria was a distant second, and detritus (coprophagy) was a negligible fraction.
Further studies of this component are required and e!orts to correct the paucity of
zooplankton data in this region are currently underway (Hopcroft et al., 1999;
GLOBEC report 17, 1996).
5.5. General conclusions
The model reproduces the observed integrated primary production, phytoplankton
biomass and nitrate concentrations within the mixed layer of the coastal upwelling
system of Monterey Bay. The model also estimates the #ows between the compartments of the planktonic ecosystem. The #ows describe the dynamics of the system
better than biomass, and facilitate comparison with other ecosystems (Ulanowicz,
1986). The results of the model con"rm the major role of upwelling as a physical force
driving the primary production cycle. This role is not limited to setting the magnitude
and pattern of nitrate supply, but is also important in terms of horizontal advection of
nutrients and phytoplankton out of the system.
The success of our modeling e!ort is due in part to the relative wealth of measured
input and validation parameters available for the region. Time series, like those at
Bermuda, Hawaii and Monterey Bay, are recognized as valuable when they are

1102

R.A. Olivieri, F.P. Chavez / Deep-Sea Research II 47 (2000) 1077}1106

needed, but on a day-to-day basis often appear routine. These e!orts are resource
intensive and hence they are often ignored or given low priority for funding. However,
they provide essential background information for process studies, are needed for
detecting seasonal, interannual and long term trends, and for modeling.
Finally, ecosystem models will be needed to explore relationships between longterm variations in ocean physics (White et al., 1997), biogeochemical cycling and
"sheries (GLOBEC report 17, 1996). A logical step is to incorporate an NPZ model
into the improving three-dimensional physical models of the northeast Paci"c. The
MB model (or a derivative), embedded in a general circulation model of the northeast
Paci"c, could be used to investigate El Nin8 o e!ects, the dramatic ecosystem changes
that have taken place in the mid-1970's (Roemmich and McGowan, 1995) as well as
the response of the ecosystem to climate change.

Acknowledgements
We gratefully acknowledge the "nancial support of the David and Lucile Packard
Foundation. Reiko Michisaki and Steve Lowder provided exceptional technical
support.

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