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James Fay 23415045

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Using an NPZD Model to Evaluate HNLC Dynamics at Kerfix Station within
the Southern Ocean, 5040'S, 6825'E
High Nutrient Low Chlorophyll regions are characterised by low phytoplankton and
high nutrient levels. By using a simple NPZD model and assessing key parameters such
as phytoplankton production rates, zooplankton grazing rates, assimilation efficiency
and mortality rates the effects they have upon HNLC regions can be observed and
documented. From the proposed model it was found that zooplankton grazing rates
play a key role in controlling HNLC zones as they control phytoplankton populations
through top down control. From the model, interactions between the phytoplankton and
zooplankton are of significance to maintaining HNLC states regardless of nutrient
limitation which according to the model has less influence on maintaining these HNLC
regions than interactions between the zooplankton and phytoplankton.
Introduction
High nutrient low chlorophyll (HNLC) regions are areas within the ocean where there is an
abundant supply of nutrients available for the use by primary producers, however despite this,
phytoplankton concentrations remain low (Edwards et al. 2004). Due to this unusual
phenomenon, it has sparked scientific interest as to why do phytoplankton remain in such low
concentrations despite high nutrient availability, where usually phytoplankton levels would
strip the system of nutrients in high abundances. HNLC areas are usually found at lower
latitudes within the Southern Ocean (Fujii et al. 2005), where chlorophyll concentrations are
low, which is shown within (figure 1). From this it could be said that HNLC regions are
latitudinal in distribution.

Fig1. SeaWiFS satellite image of chlorophyll content, green and yellow indicates high
content whereas blue and purple show low content (Feldman, 2000).
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The Study of HNLC regions is an important topic as phytoplankton play a major role within
the cycling of nutrients, cycling up to 50% of photosynthetic carbon, which within a climate
of increasing CO
2,
which is causing increased global warming is becoming more important.
By understanding what is limiting phytoplankton growth potential measures may be put in
place, in an effort to increase their concentrations within these regions which could act as a
major sink of atmospheric carbon from the atmosphere to deep oceans (Moroney & Ynalvez,
2009). The use of models in learning about HNLC processes has been claimed to be a vital
study tool of these areas (Boyd et al. 2000). This can be used to help understand how
different processes impact on these regions. This study is vital to understand consequences of
any CO
2
management protocols that may be put in place within the future, which may have
detrimental effects to the ecosystems of the oceans.
The aim of this simple model presented is to replicate conditions that occur within the HNLC
region found at the Kerfix station located at 5040'S, 6825'E within the Southern Ocean.
Where the model will be fitted to data collected, from there sensitive parameters will be
changed in order to assess their impacts in controlling and maintaining the HNLC state found
here.
Various hypotheses have been put forward as to what is controlling these HNLC regions
including: grazing rates, growth rates, mortality rates, light limitation and iron limitation
(Mitchell et al. 1991; Fasham, 1995; Pondaven, 2000). Within the model these hypothesis
will be tested to see what effects they have upon the HNLC region at Kerfix excluding iron
which its concentration is not considered within the model.

Model Description and Methods
The model used within this analysis of HNLC systems is represented by the flow diagram
within (figure 2), when run the model is sensitive to initial conditions and output will change
as a result of differing values. The model is split into four compartments, which consider:
nitrate (N), phytoplankton (P), zooplankton (Z) and detritus (D). Varying parameters control
the interactions between these compartments which are listed within (table 1) and how they
interact with each other is shown within the model equations and flow diagram (figure 2).
The model is also less complex compared to previous studies as it does not consider bacteria,
which have been shown to have a small impact on the model as a result it will not be
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considered within this model (Pondaven, 2000). Furthermore the model also
compartmentalises grazing into one zooplankton term and does not consider differing life
cycles and feeding strategies of herbivores and carnivores. It also groups dissolved inorganic
nitrogen (DIN) as one term and does not consider ammonium, thus the model follows a four
compartment model, instead of a 6 compartment model making it less complex (Fasham,
1995).

Fig2. Flow diagram illustrating the pathways of the model of how HNLC processes occur
and how N may be cycled.

Nitrate
Phytoplankton
Detritus
Zooplankton
Surface
Ocean
Deep Ocean
deepN
Primary production
Phytoplankton
mixing
Nitrate
mixing
Phytoplankton
Detrainment
Nitrate
Detrainment
Phytoplankton
Uptake, Q
Grazing
Messy feeding
(pellets)
Phytoplankton
mortality
Detrital
Remineralisation
Zooplankton
detrainment
Detrital
Mixing
Detrital
Detrainment
J
Zooplankton
Mortality
MLD
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Parameters &
Variables
Description Standard Value
N Nitrate, mmol N m
-3
30
P Phytoplankton, mmol N m
-3
0.2
Z Zooplankton, mmol N m
-3
0.1
D Detritus, mmol N m
-3
0.1
k
mix
mixing at base of mixed layer, d
-1
0.1
d
N
nitrate below mixed layer, mmol N m
-3
30
P max. phytoplankton growth rate, d
-1
2
Pm max. rate photosynthesis, d
-1
2
initial slope of P-I curve, (W m-2)
-1
d
-1
0.025
mP phytoplankton mortality rate, d
-1
0.015
kN half sat. const. for nut. Uptake, mmol N m
-3
0.5
g max. zooplankton grazing rate, d
-1
2
kg half sat. const. for grazing, mmol N m
-3
1
n shape parameter for functional response 2
zooplankton assimilation efficiency 0.75
fraction zooplankton loss term to detritus 0.25
mZ zooplankton max. mortality rate, d
-1
0.25
kZ zoo. half sat. const. for loss, mmol N m
-3
1
mD detrital remineralisation rate, d
-1
0.05
VD detrital sinking rate, m d
-1
10
kw light attenuation by water, m
-1
0.04
kc light attenuation by phyto, m
2
(mmol N)
-1
0.03
parfrac fraction of total irradiance that is PAR 0.41
latitude latitude, degrees -50
solarconst solar constant, W m
-2
1,368
albedo ocean albedo (reflectance) 0.06
cloud cloud fraction, oktas 6
Table1. List of the parameters and variables with initial standard values which represent a
HNLC system.
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The initial values for each parameter were chosen based on previous SOES 6021 practicals
which also looked at the Kerfix station and from the values provided within the model
provided by the assessment folder. A sensitivity analysis was performed using these values in
order to find the parameters that the model is sensitive to (figure 3 & 4). Initial parameter
values were then obtained by altering these parameters so that the model fitted the data
accurately. The changed values were then checked using literature to ensure that they were
realistic and could occur in situ (Anderson & Nival, 1989; Frost, 1991; Fasham, 1995).
Model Equations
The model constructed in (figure 2) was constructed using the equations below using
parameters within (table 1).

dt
dN
=
|
|
.
|

\
|
+
+
O D m
Z Z k
Z m
D
Z
Z
) (
) 1 ( +
( )
MLD
N d
N

mix
k
- ) (N H - P Q J
P
.

The nitrate compartment is represented by the above equation; nitrate is supplied by and
replenished by the cycling of dead zooplankton and remineralisation of detrital matter. It is
also further supplied by mixing with nitrate from the deep at the mixed layer depth (MLD).
Nitrate is lost from the system by detrainment to the deep ) (N H and from uptake by
phytoplankton which is used within photosynthesis and phytoplankton growth P Q J
P
. .
where J is the amount of light that is available to the phytoplankton (Evans & Parslow, 1985);
Q is the uptake rate of N and P
P
is the phytoplankton growth rate.

dt
dP
= P Q J
P
. -
MLD
P k
mix
- ) (P H - GP - P m
P

Z P kg
gP
GP
n n
n
) ( +
=

The phytoplankton compartment is influenced by the phytoplanktons ability to grow
P Q J
P
. which is described within the nitrate compartment equation description. It is then
exploited by grazing pressure forced upon by zooplankton which is represented as GPwhich
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considers zooplanktons feeding strategy and displays Holling-type III behaviour. Other loss
terms include the mortality rate of the phytoplankton P m
P
and loss of phytoplankton via
mixing at the MLD and detrainment to the deep ocean ) (P H .

dt
dZ
= GP . | - ) (Z H -
Z Z k
Z m
Z
Z
) ( +

Z Z k
Z m
Z
Z
Z
mort
) ( +
=

Zooplankton processes are represented by the above equation where the zooplanktons growth
depends on its grazing of phytoplankton, GP and the assimilation efficiency of this | . Where
the unassimilated food is lost to D. Zooplankton is lost via mortality represented by the Z
mort

equation and detrainment to the deep ocean ) (Z H .

dt
dD
= GP ) 1 ( | + P m
P
+
|
|
.
|

\
|
+
O
Z Z k
Z m
Z
Z
) (
-
MLD
D k
mix
- D m
D
-
MLD
D VD.
- ) (D H

The detritus component of the model has multiple inputs from, unassimilated food from
zooplankton in the form of pellets, GP ) 1 ( | , dead phytoplankton that is cycled to D, P m
P
,
and a fraction of dead zooplankton, , that is cycled to D. Not all zooplankton are lost this
way as some are most likely predated upon and passed up through the food chain. Loss terms
from detritus include mixing at the MLD, the remineralisation rate of detrital particles, D m
D
,
the sinking rate of detrital particles, VD and the detrainment of D via ) (D H .
The model is also influenced by physical forcing factors that are characteristic of the Kerfix
stations geographic location. These were used to control seasonal inputs of the model. The
factors that were considered were the daily clock and how much sunlight is available each
day, the irradiance that is present at this latitude and the depth of the mixed layer depth.

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Results
The models initial starting parameter values (table 1) were obtained by first running a
sensitivity analysis upon the biological parameters of the model in order to see which
parameters the model was sensitive to and which parameters affected the Phytoplankton max
(figure 3) and Nitrate max (figure 4) values. These values look at phytoplankton and nitrate
concentrations and using sensitivity analysis the parameters can be changed in order to fine
tune the models output values close to data values collected from the Kerfix station HNLC
area (figure 5). Other parameters were not considered during the sensitivity analysis as these
are unlikely to be the cause of maintaining HNLC areas as hypothesised.


Fig3. The sensitivity analysis of biological parameters affecting Phytoplankton max (Pmax).
Pmax Sensitivtiy
Biological Parameters
up Pm alpha mP g kg ngraz beta omega mZ kZ
S
e
n
s
i
t
i
v
i
t
y

t
o

P
m
a
x
-2.0
-1.5
-1.0
-0.5
0.0
0.5
1.0
1.5
2.0
+10%
-10%
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Fig4. The sensitivity analysis of biological parameters affecting the Nitrate max (Nmax).
After running the model using the initial parameters within (table 1) the output can be seen
within (figure 5). Of which the model run fits closely to the data obtained at Kerfix. Within
the phytoplankton concentration of (figure 5) the high phytoplankton data was assumed to not
be part of normal HNLC features as it characterises high phytoplankton concentrations, as a
result it is most likely a mesoscale feature passing through which can cause high
phytoplankton levels (Dagg et al. 2006). As the aim of this model is to assess the effects that
control HNLC states the model was tuned to fit the lower concentration levels which are
likely more representative of the HNLC system. Furthermore due to the close fit the model
produces to the data, the initial values could be said to be close to what may actually be
occurring at the Kerfix station.
Nmax Sensitivity
Biological Parameters
up Pm alpha mP g kg ngraz beta omega mZ kZ
S
e
n
s
i
t
i
v
i
t
y

t
o

N
m
a
x
-0.20
-0.15
-0.10
-0.05
0.00
0.05
0.10
0.15
0.20
+10%
-10%
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Fig5. Model output using standard parameter values where Nutrients (top left) and
Phytoplankton (top right) are shown at a HNLC state. Predicted PZD values during the run of
the model are shown (bottom left) and N (bottom right).
Simple Destabilising effects on HNLC State

Fig6. Raising the max phytoplankton growth rate from 2 d
-1
to 4 d
-1
Nutrient (left)
phytoplankton (right) all other parameters are standard value.
0 500 1,000
0.0
0.5
1.0
P
Z
D
day
P
,

Z
,

D
Non-commercial use only!
0 500 1,000
0
5
10
15
20
25
30
day
N
Non-commercial use only!
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10


Changing the phytoplanktons growth rate increases the amount of phytoplankton that grows
thus theoretically their abundance should increase, however in (figure 6) there is only a slight
increase in phytoplankton concentrations but there is a large decrease within the nutrient data.
This means that the increased growth rate has caused an increase in the phytoplankton but
they are immediately being removed from the system possibly via top down control.

Fig7. Raising the Phytoplankton-Irradiance from 0.025 (W m-2)
-1
d
-1
to 0.14 (W m-2)
-1
d
-1
as
seen in (Fasham, 1995) Nutrient (left) phytoplankton (right) all other parameters are standard
value.
Raising the phytoplankton-irradiance value (figure 7) allows for more efficient
photosynthesis, thus increasing primary production, which allows for increased
phytoplankton concentrations however like (figure 6) the phytoplankton concentrations are
still being suppressed. This could be top down control.

Fig8. Lowering the Zooplankton grazing rate from 2 d
-1
to 0 d
-1
Nutrient (left) phytoplankton
(right) all other parameters are standard value.
0 500 1,000
0
5
10
15
20
25
30
P
Z
D
N
day
P
,

Z
,

D
Non-commercial use only!
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Within (figure 6 & 7) increasing the phytoplankton growth rate certainly has the potential to
deplete nutrients and increase phytoplankton abundances and destabilise the HNLC system.
However these rates are being controlled by another variable which is maintaining the HNLC
state, within (figure 8) it shows the potential controlling factors limiting the phytoplankton
growth. By removing the grazing pressure, the phytoplankton are allowed to grow
uncontrolled where they grow to large concentrations, 16 mmol N m
-3
, until nutrients become
depleted where the phytoplankton concentrations then collapse, thus the limiting factor
becomes nutrient availability.

Fig9. Lowering the assimilation efficiency of zooplankton from 0.75 to 0.25 Nutrient (left)
phytoplankton (right) all other parameters are standard value.
By lowering the assimilation efficiency of the zooplankton ( figure 9) it causes slower growth
of the zooplankton population, as a result they are unable to respond quickly to increasing
phytoplankton concentrations, this allows for a higher phytoplankton concentration which
depletes nutrient concentrations, this could also be a contributing factor to the suppressing of
phytoplankton effect seen in (figure 6 & 7).
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Fig10. Raising the zooplankton mortality from 0.25 d
-1
to 2 d
-1
Nutrient (left) phytoplankton
(right) all other parameters are standard value.
Increasing the mortality of the zooplankton causes less zooplankton to be present within the
system; this means that there is less grazing pressure exerted upon the phytoplankton
population. This allows for increased phytoplankton concentrations and decreased nutrient
concentrations. Changing the mortality has a similar effect to changing the assimilation
efficiency and grazing rates.
From (figures 6 to 10) changing the parameters in one direction has opposite effects to the
HNLC state. For example the above figures show that changing the parameters in the
proposed way has destabilising effects to the HNLC system. However by changing them the
other way i.e. decreasing the zooplankton grazing rate will cause the HNLC system to
become more stable as phytoplankton concentrations are lowered and nutrients increased.







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Complex effects on a HNLC

Fig 11. Changing the rate of photosynthesis (top row) increasing photosynthesis from 2 d
-1
to
5 d
-1
, (bottom row) decreasing photosynthesis from 2 d
-1
to 0.1 d
-1
, Nutrients(left),
phytoplankton (middle), predicted P, Z values during model run (right).
Interestingly raising the rate of photosynthesis (figure 11) has a stabilising effect on HNLC
system and helps to maintain it. Whereas lowering the photosynthesis rate destabilises the
HNLC. This could be said to be unexpected as raising the rate of photosynthesis should allow
faster phytoplankton growth thus allowing for a higher abundance to occur, however the
opposite occurs, which is due to stronger coupling with the zooplankton as the increased rate
of photosynthesis allows for rapid blooming events to occur causing phytoplankton
populations to increase rapidly, as a result the zooplankton also increase rapidly and deplete
the phytoplankton quickly. Due to this they have less of an impact on the nutrients. On the
other hand lowering the photosynthesis rate causes weaker coupling with zooplankton as the
phytoplankton bloom slower, which means it takes longer for the zooplankton population to
react to the changing phytoplankton concentrations. As a result the phytoplankton bloom
persists for a longer period of time, thus having a bigger impact upon the nutrient
concentrations. This effect can be seen in more detail by removing the grazing impact of
zooplankton within (figure 12).
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Fig12. Removing the grazing pressure of zooplankton, (top) increased photosynthesis to 5d
-1
,
(bottom) decreased photosynthesis to 0.1d
-1
Nutrient (left) phytoplankton (right).
By removing grazing, nutrient concentrations become the limiting factor, where the
photosynthesis rate is increased phytoplankton concentrations are allowed to attain high
concentrations. However, they deplete the nutrients quickly and therefore collapse quickly
allowing nutrients to recover. On the other hand lowering the photosynthesis rate limits the
maximum growth rate of the phytoplankton as a result their concentrations are lower, but as a
result they deplete nutrients slower allowing for their concentration to persist for longer at
high concentrations, causing a larger impact on the nutrient concentrations.
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Fig13. Changing the phytoplankton mortality rate from 0.015d
-1
to (top) mortality rate = 0d
-1
(bottom) mortality =0.1
-1
(left) nutrient, (bottom middle), phytoplankton, (top right)
phytoplankton, (bottom right) predicted P, Z, D values during model.
With complete removal of mortality (figure 13), the HNLC is destabilised slightly, yet is still
a HNLC state as the phytoplankton are still being removed from the system by grazing. In
addition raising the mortality helps to stabilise the HNLC as phytoplankton are removed from
the system more quickly. As a result of this mortality has no destabilising effect upon the
HNLC as its effect has been replaced by zooplankton grazing, which indicates that
zooplankton grazing is a bigger loss term of the phytoplankton than their mortality rate.

Discussion
By assessing the sensitive parameters of the model that affect the state of HNLC systems, it
has therefore been possible to then test the effects of these parameters upon a HNLC system,
in order to find out which parameters are controlling this phenomenon from occurring. The
parameters that were found to be sensitive to the model were also the parameters put forward
within the hypothesis.
0 500 1,000
0.0
0.5
1.0
P
Z
D
day
P
,

Z
,

D
Non-commercial use only!
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Of the parameters that were tested the grazing rate of zooplankton was found to be one of the
most dominating processes (Martin et al. 1990; Fasham, 1995; Sato et al. 2007) and is an
important process in regulating phytoplankton populations. This was evident as although
increasing the growth and light available to the phytoplankton their populations were still at
low concentrations; which is most likely due to the top-down control from zooplankton
(Landry et al. 2011; Adjou et al. 2012). As zooplankton is found to be one of the biggest
factors influencing phytoplankton populations via grazing, changing parameters that
influence the grazing rate are the next biggest factor. This was seen within the results as the
removal of grazing allowed for explosive and unchecked growth of the phytoplankton until
they became nutrient limited. In addition by raising mortality of zooplankton and lowering
the assimilation efficiency, it lowers the growth of the zooplankton population, which results
in a lower grazing rate through reduced abundances, due to this, these factors are the next
biggest factor controlling the HNLC state (Landry et al. 2011; Adjou et al. 2012).
As the grazing rate of zooplankton upon phytoplankton is a key loss term for phytoplankton
and a main controlling factor of phytoplankton death, the phytoplankton mortality is therefore
mainly controlled by the grazing pressure. As a result the natural phytoplankton death
through other means than grazing does not have a significant effect as grazing replaces it as
the main factor causing phytoplankton mortality (Mitchell et al. 1991; Landry et al. 2011;
Adjou et al. 2012). Therefore if you increase grazing rates then natural mortality could be
expected to decrease as a result.
Therefore due to grazing pressure and the top down control from zooplankton, parameters
that affect phytoplankton growth such as growth rate, light availability and photosynthesis
rate are secondary factors that control HNLC regions which is observed in the results; as
where grazing is present populations are still low despite increasing the production of
phytoplankton which is due to grazing. However, by removing the grazing factor the
phytoplankton production factors then become major controls of HNLC regions (Mitchell et
al. 1991; Fasham, 1995; Moroney & Ynalvez, 2009).
To conclude interactions between the phytoplankton and zooplankton have significant effects
on causing HNLC states to occur and community level interactions play an important role in
maintaining HNLC regions (Landry et al. 2011).
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The last major controlling feature upon HNLC regions is the nutrient concentrations, as
despite their high abundances they are not being used, this is because grazing rate suppresses
phytoplankton concentrations which draw down nutrient concentrations. As a result if grazing
is removed and phytoplankton growth rates are sufficient then nutrients then become the
limiting factor, which can be seen within (figure 8). It is therefore unlikely that the iron
hypothesis is the exclusive reason HNLC regions occur and is more likely controlled by the
zooplankton coupling with phytoplankton; in addition zooplankton grazing has been shown
to be an important process in the remineralisation of iron (Fasham, 1995; Boyd, 2002; Sato et
al. 2007). Having said this due to phytoplankton being assumed as one compartment this
model is unable to distinguish between plankton that require iron and those that do not, as a
result it not possible to draw credible conclusions about the effects of iron on the
phytoplankton using this model. Furthermore considering that most models now take this
factor into account and has shown that there is a significant difference between these groups
of phytoplankton, there is a need for a more complex model in order to assess the impact of
iron (Hinckley et al. 2009).
By using models to understand HNLC regions it is possible to understand the level of
influence a process may be having on the system, in doing so it is possible to find out which
process is the main cause for the occurring phenomena. Therefore the use of models is
important in mapping out the role of parameters upon a system (Boyd et al. 2000). However
it is important to take into consideration the limitations of the model that is being used as
simplicity of a model will overlook many potential key processes. In addition the level of
knowledge available for constructing a complex model is limited as the life cycles of many
key phyla is limited or unknown. This forces compartmentalising which can increase error
within the model (Fasham, 1995). Additional errors for models is that they assume a closed
system and are not programmed to handle external influences such as mesoscale features
which may pass through the system these can influence the results and as seen with the above
data set could cause doubt with the models validity (Dagg et al. 2006).

References
Adjou, M., Bendtsen, J., Richardson, K. (2012). Modeling the influence from ocean transport,
mixing and grazing on phytoplankton diversity. Ecological Modelling, 225, 19-27.
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Anderson, V., Nival, P. (1989). Modelling of phytoplankton population dynamics in an
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