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Aquaculture pond ecosystem model: Temperature and dissolved oxygen

prediction - Mechanism and application

Article  in  Ecological Modelling · August 1996

DOI: 10.1016/0304-3800(95)00140-9

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 E[OLO61I:IIL
mOOELLIn6
ELSEVIER Ecological Modelling 89 (1996) 231-258

Aquaculture pond ecosystem model: temperature and dissolved

oxygen prediction -
mechanism and application
Steven D. Culberson, Raul H. Piedrahita *

Department of Biological and Agricultural Engineering, University of California, Davis, CA 95616, USA
Received 15 August 1994; accepted 26 April 1995

Abstract

Accurate characterization of temperature and dissolved oxygen stratification in ponds used for aquaculture is of
critical importance in understanding how these ponds may be constructed, oriented, or otherwise managed
biophysically when one wishes to provide optimal environmental conditions for the organisms being cultured. While
field studies can provide characterizations of water quality stratification at a single locale, to date there have been
few attempts at developing reliable models which can be used at a variety of sites after initialization with appropriate
local geographic and atmospheric data. In conjunction with the Pond Dynamics/Aquaculture Collaborative Re-
search Support Program ( P D / A CRSP), the authors have developed a water quality model (the Aquaculture Pond
Ecosystem Model, A.P.E.M.) which predicts temperature and dissolved oxygen stratification at several sites,
including Thailand, Rwanda, and Honduras. Advances in model structure and reduction of data requirements
relative to previous models reflect the desire to provide for culturists the ability to predict stratification events with
commonly available data, obtained either by hand or from a simple weather station located at or near the pond site.
Strategies used for dealing with the reduced data sets, and the associated assumptions made, are also presented.
Lastly, a series of simulation runs is performed to assess the quantitative effects on temperature and dissolved
oxygen concentration generated by varying pond depth and phytoplankton density input values.

Keywords: Oxygen; Pond; Stratification; Temperature

I. Introduction of the culture environment itself - water quality

r e q u i r e m e n t s for the cultured species must be
T h e science of aquaculture d e m a n d s a wide
met and m a i n t a i n e d (Allan and Maguire, 1991;
range of skills and knowledge on the part of the
Bejda et al., 1992). A q u a c u l t u r e is carried out in
culturist, particularly as the culture system is
a wide variety of systems, and ponds are the most
p u s h e d to contain higher densities of animal and
common p r o d u c t i o n systems on a worldwide ba-
plant biomass and to p r o d u c e increasing quanti-
sis. A q u a c u l t u r e ponds are normally less than 1.5
ties of high quality product. P a r a m o u n t to this
m deep, and vary in size from u n d e r 100 m 2 to
science is an accurate and reliable u n d e r s t a n d i n g
over 10 ha. Animals cultured in these ponds may
be fed p r e p a r e d diets, or the ponds may be
* Corresponding author. Fax: ( + 1-916) 752-2640. fertilized to p r o m o t e the production of food for

0304-3800/96/$15.00 © 1996 Elsevier Science B.V. All rights reserved

SSDI 0 3 0 4 - 3 8 0 0 ( 9 5 ) 0 0 1 4 0 - 9
232 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
the fish within the pond. Regardless of how the
animals in a pond are fed, water quality is largely
determined by biological and chemical processes
related to photosynthesis and respiration by or-
ganisms within a pond. Predicting and managing
water quality in pond aquaculture presents spe-
cial challenges due to the exposure of the system
to factors of climate and topography. Since bio-
chemical processes and organism growth rates
are regulated to a large extent by temperature,
and since the survival of the organisms within the
pond environment depends principally upon the
amount of available oxygen, pond water tempera-
tures and dissolved oxygen (D.O.) levels are likely
to be the most important items of concern to
pond water quality management (Boyd et al.,
1978; Boyd, 1990).
Inasmuch as animal mortality continues to oc-
cur within the industry due to low D.O. incidents
in rearing ponds, further understanding of D.O.
dynamics within these ponds is essential, particu-
larly if this understanding leads to qualitative
a n d / o r quantitative predictions of what these
levels might be under given local atmospheric
conditions. Since the practice of mechanically
aerating ponds during periods of low D.O. is
dependent upon supplying enough oxygen to ful-
fill the respiratory needs of the animals and plants
within the pond, and since there may be some
time lag between when the aerators begin work-
ing and when the oxygen supplied is fully mixed
into the pond, prediction of the time of onset of
low D.O. events would allow more efficient appli-
cation of aerator usage.
Shallow aquaculture ponds typically show daily
patterns of temperature and oxygen stratification,
rendering portions of these ponds unusable by
the cultured animals. Stratification events ob-
served in aquaculture ponds tend to be short
lived, as ponds normally stratify during the day
and destratify at night (Chang and Ouyang, 1988;
Losordo, 1988). In ponds showing marked stratifi-
cation, surface waters may be harmful to fish due
to supersaturated D.O. conditions in combination
with high temperatures, while in the same pond
near anoxic conditions may exist close to the
bottom (Krom et al., 1985; Chang and Ouyang,
1988; Losordo, 1988; Boyd, 1990).
Previous attempts to characterize temperature
and dissolved oxygen patterns in ponds used for
aquaculture are not common in the literature,
and where they exist, most tend to be made using
direct observations of the pond in question, use-
ful only for predictions of nighttime D.O. decline,
a n d / o r are based on the assumption the pond
water column is uniform over depth (Boyd et al.,
1978; Romaire and Boyd, 1979; Meyer and Brune,
1982; Klemetson and Rogers, 1985; Madenjian et
al., 1987). There are, however, several reports of
models incorporating mechanistic characteriza-
tion of the biochemical and physical processes in
a pond which together govern the resulting tem-
perature and D.O. levels (Cathcart and Wheaton,
1987; Losordo, 1988; Losordo and Piedrahita,
1991). While fairly accurate, these models are not
very "user friendly", and require large, complete,
and complex data sets for execution.
The purpose of the study described below was
to construct a mechanistic model (the Aquacul-
ture Pond Ecosystem Model, A.P.E.M.) which
capitalized on the understanding of the biologi-
cal, chemical, and physical processes embodied in
the more complex models cited above, while sim-
plifying the data input needs so the resulting new
model would be applicable to a broader range of
shallow pond aquaculture sites where access to
data is more limited. This study has updated
information contained within the previous mod-
els, taking into account more recent characteriza-
tions of the biological and chemical processes
occurring in the pond ecosystem. The model pre-
sented in this paper was developed using a struc-
ture similar to a previous model (Losordo, 1988;
Losordo and Piedrahita, 1991), with considerable
revision. It is hoped this revision and simplifica-
tion will provide a predictive tool for farmers
a n d / o r for facility planners who may want to
know how pond morphology, siting, and manage-
ment techniques will affect temperature and dis-
solved oxygen levels in any ponds built on a
prospective site.
The current model has been validated using
data from the Pond Dynamics/Aquaculture Col-
laborative Research Support Program database
( P D / A CRSP, 1992), and this paper demon-
strates the accuracy of the temperature and D.O.
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258 233

predictions at a variety of aquaculture sites. A
subsequent discussion is presented detailing the
response of the coupled temperature and dis-
solved oxygen model to various alterations of
pond depth and phytoplankton concentration.
2. Temperature
A conceptual view of the pond water column
as it is assumed to be for the purposes of the
present model is presented in Fig. 1 (after
Losordo et al., 1986). For the purpose of charac-
terizing water column stratification, this concep-
tual diagram has been chosen to have three water
"volume elements" (termed "surface", "middle",
and "bottom"), and one s e d i m e n t / b e n t h i c vol-
ume element. The thickness of each of the vol-
ume elements will range from 0.3 to 0.5 m for
typical aquaculture ponds of depth 0.9 to 1.5 m.
It is the characterization of each of the processes
depicted in Fig. 1 which forms the bulk of the
working model. In addition, there are subroutines
for calculating solar incidence angles, effective

EnergyRux
Oxygen Pond/Atmosphere Evaporative
_ElementDep~
PondSurface z0

"~10xygenConsumption/Production ~" zl
PenetratingSolar EnergyTransfer
RadiatioIn Surface/Middle
I Effective
ConvectiveI diffusion z2
Mixing

" •OxygenConsumpti
P o1 n/Production~ z3

eI
Convectiv
Radiationl
PenetratingSolar
Energy
Mi TransferI
ddle/Bottom Effective
diffusion z4
Mixing

- •
OxygenGonsump'don/Produ~onJ~ 26

Penet~ngSolar
Radiation![ J, Sediment
26

27

z8
EnergyExchangeI
w/Groundwater

Fig. 1. Conceptual view of a stratified aquaculture pond model showing heat and mass flows (after Losordo et al., 1986).
234 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
NetEnergySurf
Idb'~ tl Ik~l a U ~ e ~ d L ~ ! I
Fig. 2. STELLA I I Q diagram of energy-balance structure.
wind fetch distances, etc., which can be thought
of as "supporting" calculations - important in
the execution and integration of all of the various
model equations, but which are not calculations
directly responsible for changes in t e m p e r a t u r e
or dissolved oxygen levels.
An assembly of the volume elements is con-
structed in the vertical plane to account for dif-
ferences in rate processes, diffusion, density, light
penetration, etc., with depth in the simulated
pond ecosystem. In all cases there is an assumed
horizontal homogeneity within any given vertical
volume element. This assumption is consistent
with data presented in studies of vertical and
horizontal t e m p e r a t u r e variation in ponds used
for aquaculture (Cathcart and Wheaton, 1987;
Losordo, 1988; Losordo and Piedrahita, 1991).
W h e n all the energy and mass balance pro-
cesses important within the conceptual diagram
are identified, the model's basic framework is
constructed using S T E L L A IIl(High Performance
Systems, Inc., 1990). Fig. 2 presents a schematic
diagram for those elements used to calculate the
energy balance in the water column. For the
purposes of the present model, the following form
of the net energy change in the pond ecosystem is
used (WRE, 1968; TVA, 1972):
~net =~)sn-{-~)at--~ws--(j~e~t-~Dc--(l~sn,z-{'-~sed
- ~ -+~d,: (1)
where:
~net = net heat flux for the pond
~'s. = penetrating short-wave solar irradiance
(Eq. A1)
q~at = net atmospheric radiation (Eq. A2)
~bws = water surface back radiation (Eq. A3)
~be = evaporative heat transfer (Eq. 2)
the = sensible heat transfer (Eq. A4)
~b.... = solar irradiance at lower volume ele-
ment boundary (Eq. A5)
~bsed = heat transfer between sediment and bot-
tom water volume elements (Eq. A6)
~bgw = heat loss from sediment volume element
to ground water table (Eq. A7)
~bd,z = effective diffusion of heat at upper and
lower volume element boundaries (Eq. A8)
N.B. All heat flux terms in kJ/mZ/hr.
In this generalized form of the heat flux equa-
tion not all terms will be considered for each of
the volume elements. T h e r e will be no ~bgw term
for the surface or middle volume elements, for
example, and there will be no ~be term for the
middle or bottom volume elements. Each of these
heat flux elements and their mathematical de-
scriptions as implemented in the current model,
are shown in Appendix A. Revisions to previ-
ously-used formulations (Losordo, 1988; Losordo
and Piedrahita, 1991) are described below.
2.1. Evaporative heat transfer
H e a t loss by evaporation from the pond sur-
face involves a state change of water from liquid
to water vapor. The energy required for this
transformation through the latent heat of vapor-
ization is taken from the energy stored within the
water column (Moore et al., 1978; Orlob, 1981).
This process is also influenced by the effects of
wind across the surface of the pond, and by the
moisture content in the air lying just above the
water surface (Henderson-Sellers, 1984). Losordo
and Piedrahita (1991) successfully employed the
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
following estimation of evaporative heat flux fol-
lowing studies by Fritz et al. (1980):
qbe = N W 2 ( e s - e a )
where:
N = empirical coefficient from Lake Hefner
(Fritz et al., 1980)=5.0593 (kJ m -2 km -1
m m H g - 1)
W2 = w i n d speed 2 m above water surface
(km/hr)
e S = saturated vapor pressure at Twc (mmHg)
e a = water vapor pressure above the pond sur-
face (mmHg)
In turn, e s and e a are approximated as (Fritz
et al., 1980):
e s = 25.374 exp(17.62 - 5271/Twc )
e a = Rh* 25.374 exp(17.62 - 5271/T~c )
where Rh = relative humidity measured 2.0 m
above the water surface.
Relative humidity measurements were not
available from the P D / A CRSP Database ( P D / A
CRSP, 1992). Henderson-Sellers (1984) offers a
slightly different estimation for water vapor pres-
sure which uses dew-point temperature in place
of relative humidity measurements:
e a = 610.78 exp[17.2694* T a ~ -273.16
-~- ]1
where Td = dew-point temperature in K.
While dew-point temperatures are rarely
recorded for aquaculture pond sites, Koon et al.
(1986) suggest average daily dew-point tempera-
ture may be estimated by subtracting 2°C from
the morning minimum dry-bulb temperature. This
estimation, in combination with Eq. 5 shown
above, is then incorporated into the present model
to estimate evaporative heat flux. Accurate esti-
mation of the magnitude of this heat flux is
critical to an accurate simulation, as it accounts
for up to one third of the total energy loss from
the ponds studied (Culberson, 1993).
2.2. Energy distribution within the water column
The two processes responsible for energy dis-
tribution in the water column (beyond those asso-
ciated with light penetration and attenuation)
235
are: (1) that concerning the distribution of energy
due to molecular diffusivity and turbulent eddy
diffusion, commonly considered together as "ef-
(2)
fective diffusion" (dominant during warming peri-
ods, see Eq. A8); and (2) that concerning the
distribution of energy due to convective transport
(dominant during cooling periods). Each of these
processes is treated at length in Sundaram and
Rehm (1973), Chapra and Reckhow (1983), Hen-
derson-Sellers (1984), Losordo (1988), and
Losordo and Piedrahita (1991). The choices of
which estimation procedures to be included here
closely follow those used by Losordo and
Piedrahita (1991).
2.3. Energy distribution by convective mixing
(3)
(4) During cooling periods convective mixing is
the dominant mechanism for energy distribution
within the water column, buoyant instabilities oc-
cur (where cooler, denser surface waters lie above
warmer, less-dense, layers), and convective trans-
port mixes adjacent layers (Losordo, 1988;
Losordo and Piedrahita, 1991). For the purposes
of modeling this process of convective mixing the
Sundaram and Rehm (1973) method has been
incorporated here: during convective mixing
(5) events the effective diffusion coefficient between
two adjacent volume elements is allowed to as-
sume some maximum value.
The value considered as the maximum effec-
tive diffusion coefficient (Emax) depends in large
measure on the size of the time step used for
model execution (Losordo and Piedrahita, 1991),
and the present model adopts the value used by
Losordo (1988): 6000 cruZ/hr.
Implementation of the energy distribution co-
efficients described above involves the compari-
son of temperatures in adjacent volume elements
in each simulation time step. If buoyant instabili-
ties do not exist, energy movement is considered
to occur by effective diffusion. If buoyant instabil-
ities do exist, then energy movement is consid-
ered to occur by convective mixing.
2.4. Wind speed and wind uector adjustments
A number of the equations used in calculating
pond temperatures incorporate wind speeds at a
236 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
given reference height above the pond surface
(Eqs. 2, A1.2, A4 and A8.5). A simple adjustment
function was developed and used by Losordo and
Piedrahita (1991) after Pankofsky and Dutton
(1984) so that wind speed data collected from any
given height from the pond surface (up to a
maximum of 10 meters) could be used as input to
the model. In the present model this function is
included in the equation:
Wadj = W z ( 1 0 / z . . . . . . )1/7
where:
W~aj = w i n d speed adjusted for a n e m o m e t e r
height ( m / s )
Wz = wind speed measured at height z ( m / s )
Zanemom = height of a n e m o m e t e r above pond
surface (m)
In addition, wind vectors could not be calcu-
lated for the P D / A CRSP Database, since the
P D / A CRSP data collection protocols do not
include wind direction measurements. An esti-
m a t e of wind vector magnitude is made, however,
through a calibration run (see below). Once this
calibration is m a d e for a given data set with a
given anemometer height, other wind
s p e e d / a n e m o m e t e r height m e a s u r e m e n t s or in-
puts are adjusted using the same calibration fac-
tor.
2.5. Wind direction and fetch adjustments
Wind direction can have a significant effect on
the extent of wind fetch across a water body
(where pond width to length ratios are smaller or
larger than one, for example), and some account-
ing for the lack of these data must be made. D u e
to the relatively small size of the ponds simulated
for the P D / A CRSP research sites (10 × 30 m,
for example), it was assumed that in order for the
effects of the winds to be at a maximum, the
largest fetch length should be used. That is to say,
in the absence of wind direction data, the winds
were assumed to be blowing along the long axis
of the ponds. In this way, then, any effects of
wind fetch and wind shear are recognized in the
model as having their maximum possible effect.
Calibration of the model for wind vector magni-
tude subsequently corrects for wind fetch and
wind shear not being at their maximum (see be-
low).
2.6. Air temperature inputs
The data reporting protocols for the P D / A
CRSP experiment sites do not require continual
monitoring of air temperatures throughout the
(6) diurnal cycle - only the daily minimum and maxi-
m u m air temperatures are recorded. Since air
temperatures at the pond site are critical for
calculations of convective heat exchange at the
pond surface, evaporative heat loss, and atmo-
spheric radiation into the pond, an estimation of
air temperatures over the diel cycle is required.
Card et al. (1976) developed a simulation algo-
rithm which predicts air temperatures based on
sinusoids derived from long-term weather data,
initialized with minimum and maximum air tem-
perature measurements. The t e m p e r a t u r e at any
given time may be calculated:
T = Tmeanq- TempSwing* P (7)
where:
TempSwing = Tmax - Tmin (°C) (8)
Tmean = (Tmax -1- T m i n ) / 2 (°C) (9)
P=a[sin(o)ltl-4)) + b sin2(o)ltl-qb)] (10)
such that:
P = t e m p e r a t u r e profile factor
a, b = coefficients in t e m p e r a t u r e profile ex-
pression (a and b set the amplitude of the
t e m p e r a t u r e swing such that Pmax= 0.5, and
p e a k to p e a k amplitude is unity. For use in the
revised model minimum and maximum temper-
atures are assumed to occur at 6:00 am and
3:00 pm, respectively, or a = 0.4484, and b =
0.2706; see Card et al. (1976) for detailed ex-
planation)
(.o1 = angular frequency, 2~r/24 ( r a d i a n s / h r )
t I = time (in hours from midnight)
~b = phase angle (sets the profile correctly with
respect to clock time; determined by the time
at which the median t e m p e r a t u r e is expected
to occur)
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
If Tmi n o c c u r s at 6 am, and Tmax occurs at 3 p m
(the approximate times at which minimum and
maximum t e m p e r a t u r e s are expected to occur at
the P D / A CRSP research sites, respectively), the
median t e m p e r a t u r e will occur at 10:30 am and:
& = ( 1 0 . 5 / 2 4 ) .27r
or:
& = 2.7489 radians
Substituting in for P and adding 6 hours to the
simulation start time (thereby initiating the simu-
lation at 6 am as opposed to midnight) yields the
t e m p e r a t u r e estimation expression:
T = ( T m a x -I- T m i n ) / 2 q- ( T m a x - Tmin) ,0.4484
* [sin(2* 7 r / 2 4 , ( t + 6) - 2.7489)
+ 0.2706, sin(2* (2* (~-/24) * (t + 6)
-2.7489))]
The use of Eq. 12 for the prediction of air
t e m p e r a t u r e s at a given site provides one impor-
tant means of reducing the data required for
model execution. The two air t e m p e r a t u r e inputs
used for this model (minimum and maximum
readings for a given day) replace 1440 microlog-
ger-recorded air t e m p e r a t u r e readings used by
I_xgsordo and Piedrahita (1991).
2. 7. Solar radiation inputs
Since the entire energy balance description
used in the model under discussion depends on
estimations of energy flux into and out of the
system, the most useful solar radiation input mea-
surement would be in units of W / m 2. Indeed,
where these m e a s u r e m e n t s are available in the
P D / A CRSP Database, they are entered directly
into the model. However, the P D / A CRSP re-
search sites had a research emphasis on phyto-
plankton dynamics, and recorded photosyntheti-
cally active radiation (PAR) only, measured as
moles of p h o t o n s / m Z / h r . Since the p h o t o m e t e r s
measure only that radiation in the photosyntheti-
cally active range of the spectrum (approximately
420-750 nm), it is a m e a s u r e m e n t which underes-
timates total radiation received by the earth in a
particular location. For this reason, for those sites
237
where only P A R m e a s u r e m e n t s are available, an
additional calibration and correction for solar
irradiance values in W / m z is necessary.
In addition, the collection of solar radiation
data at P D / A CRSP sites is far less intensive
than that utilized by Losordo and Piedrahita
(11)
(1991). For use in the present model, solar radia-
tion readings are taken at predetermined inter-
vals (of between 2 and 4 hours) as cumulative
totals, and hourly inputs of solar radiation are
obtained by averaging the cumulative totals over
the collection interval. In real terms, where
Losordo and Piedrahita required up to 720 read-
ings (one taken every minute over the course of a
12-hour solar period) to generate inputs, sample
data sets used for the present model require as
few as five readings for data input generation.
(12) 2.8. Temperature model calculation procedure
As implemented in S T E L L A II ( ~ (High Per-
formance Systems, Inc., 1990) and determined by
the structure given to the present model, temper-
ature calculations for each of the volume ele-
ments involve tabulating the net heat flux for
each element at each time step. This net heat flux
is then added or subtracted from the previous
time step temperature, and the current time step
t e m p e r a t u r e is solved for. At the beginning of the
simulation (time = 0) the t e m p e r a t u r e of each
volume element is initialized with the 0600 hour
(dawn) measured temperature, and each
element's net heat content is set to zero. This
procedure is explained in more detail in Losordo
and Piedrahita (1991), although the calculation of
the t e m p e r a t u r e of any volume element (Vii) at
any time (t) is fairly straightforward:
Tvi,t = Tvi,o -]- ( H V i , t _ 1 -[- ( d H v i / dt ) * d t )
/ p * Cpw * v i (13)
where:
Tvi,, = t e m p e r a t u r e of Vii at time t (°C)
Tvi,o = t e m p e r a t u r e of V/ at time t = 0 (°C)
Hvi,t-a = heat stored in V// at time t - 1 (kJ)
d H v i / d t = heat flux during the time interval
(kJ/hr)
dt = duration of the time interval (hr)
238 S.D. Culberson, R.H. Piedrahita/Ecological Modelling 89 (1996) 231-258

p = density of w a t e r ( k g / m 3) however, only o n e p o n d p e r study site is pre-

Cow = heat capacity of w a t e r ( k J / k g °C)
vi = v o l u m e of v o l u m e e l e m e n t Vi (m 3)
2.9. Input data summary and sample data set
T h e d a t a collection protocols at the P D / A
C R S P r e s e a r c h sites follow a fairly u n i f o r m
cross-site p a t t e r n , particularly for those p a r a m e -
ters which describe local a t m o s p h e r i c c o n d i t i o n s
( P D / A C R S P , 1992). I n general, the data used
for execution of the c u r r e n t m o d e l were collected
at the following local times: 0600, 1000, 1400,
1600, 1800, a n d 2300 hours each day. Not every
p a r a m e t e r n e e d e d for m o d e l e x e c u t i o n was m e a -
s u r e d every day in every p o n d at a p a r t i c u l a r site,
b u t several p o n d s at each site for each of several
days were chosen for s i m u l a t i o n since a d a t a set
c o m p l e t e e n o u g h for m o d e l initialization a n d exe-
c u t i o n was available. D u e to space limitations,
s e n t e d in this report, a l t h o u g h the results de-
picted below are r e p r e s e n t a t i v e of o t h e r p o n d s
from each respective P D / A C R S P study site.
A s u m m a r y of the data i n p u t s n e e d e d for
execution of the t e m p e r a t u r e s i m u l a t i o n p o r t i o n
of the m o d e l is p r e s e n t e d in T a b l e 1 for each of
the P D / A C R S P r e s e a r c h sites. T h e b u l k of the
data i n p u t s are initialization terms. O n l y solar
r a d i a t i o n a n d wind speed i n p u t s are n e e d e d as
i n p u t variables over the course of the s i m u l a t e d
day.
2.10. Calibration for use at PD / A CRSP sites
Before the c u r r e n t m o d e l could be imple-
m e n t e d using P D / A C R S P D a t a b a s e inputs, a
c a l i b r a t i o n r u n h a d to b e executed to o b t a i n
accurate, site-specific estimates for the relation-
ship b e t w e e n w i n d speed a n d w i n d vector, a n d

Table 1
Temperature data inputs for CRSP PD/A research sites
Input variable Local time Honduras Rwanda Thailand
Julian day N/A 351 146 96
Latitude N/A 14.43°S 2.67°S 14.18°N
Pond length (m) N/A 32.0 30.0 30.0
Pond width (m) N/A 32.0 20.0 10.0
Pond depth (m) N/A 0.80 1.16 0.96
Maximum air temperature (°C) ~- 1500 36.5 28.0 37.0
Minimum air temperature (°C) ~ 0600 14.3 10.0 27,0
Anemometer height (m) N/A 3.0 2.0 2.0
Secchi disk depth (m) -~ 1200 0.13 0.52 0.35
Initial surface temperature (°C) 0600 23.0 21.0 29.8
Initial middle temperature (°C) 0600 23.2 21.0 29.8
Initial bottom temperature (°C) 0600 23.2 21.0 29.8
Solar irradiance (mols/m 2) 0600 0.00 5.01 0.00
1000 10.75 12.35 7.49
1430 18.55 5.93 22.22
1600 - 2.68 7.02
1800 6.56 0.04 2.22
Wind speed (km/hr) 0600 0.00 1.67 0.00
1000 7.39 1.67 5.50
1430 10.45 1.67 8.90
1600 - 1.67 7.40
1800 9.31 1,67 9,70
1900 9.72 - -
2000 7.94 1.39 -
2200 5.77 1.39 9.80
0600 3.97 1.39 3.10
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258 239

Measured Temperatures
for the relationship between P A R and total solar 28"
irradiance. This calibration run was conducted 27"
for a given pond at each of the three P D / A G"
0,.,
26'
CRSP research sites discussed (Honduras, P
Rwanda, and Thailand), and estimates were ob- .9.= 25'
tained for the site-specific parameters needed. 24
The calibration run-derived values were then ap- E
I.- 23
plied to different ponds at each site and simula-
tion results were generated for these novel ponds. 22
o " ; " i " ,'2 " 1"6 " 24
The need for calibration of the present tempera-
ture model would be removed if solar radiation
inputs were for total solar irradiance in addition
Simulated Temperatures
to PAR, and if in addition to wind speed, wind 28"
direction inputs were available. In all cases, the
results presented below have been generated for o~ 27'
ponds which have not received these specific cali- v
o
26'
bration treatments. 25'

2.11. Simulation results from PD / A CRSP re- 24'

E
search sites )- 23'

22
2.11.1. Honduras 0 ; 8 1'2 1'6 ='0
Data for the execution of the present model Time (0 equals 0600 local hour)

for the E1 Carao Aquaculture Experiment Station
in Comayagua, Honduras, were collected on Ju-
lian Day 351 (December 17), 1990, and the pond
simulated was #B7. Qualitatively, the timing of
maximum stratification, the remixing of the water
column after sundown, and the rate of nighttime
cooling are well represented (Fig. 3). Quantita-
tively, the model shows errors in its absolute
predictions, although even when most inaccurate
(in this case for the middle volume element), the
model under-estimates temperatures according to
a time lag function only. Even with this time lag
error, over the course of the simulation predicted
temperatures do not differ from measured tem-
peratures by more than 1.0°C.
2.11.2. Rwanda
Data for the execution of the present model
for the Rwasave Pond Dynamics/Aquaculture
CRSP Research Site at Rwasave, Rwanda, were
collected on Julian Day 146 (May 26), 1989, and
the pond simulated was #C1. The surface and
bottom layer temperatures are represented fairly
closely, but there is some disagreement between
measured and predicted middle volume element
Fig. 3. Measured and simulated temperature profiles, Hon-
duras, Julian Day 351, 1990.
temperatures (Fig. 4). This misrepresentation of
the middle layer suggests the model over-esti-
mates the amount of thermal energy diffusing
from the surface layer to the middle layer. While
significant, the error in estimation for the middle
layer does not exceed 1.5°C.
2.11.3. Thailand
Data for the execution of the present model
for the Ayutthaya Station facility at Ayutthaya,
Thailand, were collected on Julian Day 96 (April
6), 1988, and the pond simulated was #12. There
are inaccuracies for the prediction of individual
temperatures at specific times, but the overall
character of the simulation is close to what was
measured in the pond (Fig. 5). The inaccuracies
in the simulated output seem to reflect an overall
underestimation of the amount of solar radiation
throughout the 24-hour period. As with the other
simulations presented here, the timing of peak
stratification and remixing are closely predicted.
240 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258

Measured Temperatures Measured Temperatures

27" 33

p 32
25 ¸
= ~ 31
24
P 2.3'
o ~ 30
E 22' E
I--
21
20 28 , . ,% . , . .

0 4 8 1 '2 16 2'0 2'4 4 8 12 16 20 2'4

Simulated Temperatures Simulated Temperatures

27" 33'
26"
F
v 25-
Middle
P 24" = 31,

t-- 21 ~ 29

20 - ' . . . . . . . . 28
0 4 8 12 16 20 24 o ; ; t'2 l'e 2'o 2,
~me (0equals0600 local hour) Time (0 equals 0600 local hour)
Fig. 4. M e a s u r e d and s i m u l a t e d t e m p e r a t u r e profiles, Rwanda,
Fig. 5. Measured and simulated temperature profiles, Thai-
Julian Day 146, 1989.
land, Julian Day 96, 1988.

2.12. Comparison of actual and simulated tempera-

ture profiles described above, and the corresponding model
prediction of these same events. The qualitative
Table 2 presci, ta a ~ummary of the actual nature of the simulation is quite accurate; i.e. the
timing and extent of stratification in the ponds onset of stratification, the timing of the maximum

Table 2
Comparison of actual and predicted thermal stratification events
Site and Julian Stratification Maximum Isothermal Maximum
pond day onset a stratification conditions b stratification ~
Actual Simulated Actual Simulated Actual Simulated Actual Simulated
(h) (h) (h) (h) (h) (h) (°C) (°C)
Honduras
Pond # B 7 351-352 0900 1000 1500 1500 2100 1900 3.20 3.98
Rwanda
Pond # C 1 146-147 1000 1100 1600 1600 0200 0200 4.00 3.41
Thailand
Pond # 1 2 96-97 1100 1100 1600 1500 2200 2200 3.30 2.81

a T e m p e r a t u r e differences greater than 0.5°C are considered stratified.

b T e m p e r a t u r e differences less than or equal to 0.5°C are considered isothermal.
c Measured as the difference between surface and bottom readings.
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
stratification, and the resumption of isothermal
conditions in the ponds are very closely pre-
dicted. Less accurate were the prediction of indi-
vidual temperatures at each hourly interval, al-
though for the maximum stratification event the
model was within 1.0°C for all three simulated
ponds.
2.13. D&cussion: Temperature
W h a t is most striking about the model simula-
tions outlined here is the sustained accuracy of
the predictions in t e m p e r a t u r e stratification (or
lack thereof) across different pond sites in several
geographic regions. Combined with increased
flexibility of use and adaptability to less intensive
input data files, it is evident the energy-balance
approach used in these models is an appropriate
one to use when characterizing the overall heat
content within ponds used for aquaculture. The
onset of stratification, the degree of stratification
from surface to pond bottom, the onset of mixing,
and the simulation of completely mixed condi-
tions remain accurate. Differences in instanta-
neous t e m p e r a t u r e predictions from measured
t e m p e r a t u r e values across the pond depth rarely
exceed 1.0°C, and are principally associated with
a factor of time lag only.
3. Dissolved oxygen
As was the case with the t e m p e r a t u r e model
outlined above, the correct modeling of dissolved
oxygen concentrations in an aquaculture pond
depends upon an accurate description of those
factors contributing to oxygen entry into the pond,
oxygen removal from the pond, and oxygen distri-
bution within the pond. The form of the net
oxygen change in the pond water column can be
described as (Losordo, 1988):
MO2net = MO2p --I-MO2D -- M O 2 f - MO2p r
-- MO2wcr - MO2s r ___MO2d,z -- MO2x ~
(14)
241
where:
MO2net = net oxygen mass change for the vol-
ume element
MOzp = p h y t o p l a n k t o n oxygen mass produc-
tion (Eq. 15)
MOzD = oxygen mass diffusing t o / f r o m atmo-
sphere at surface (Eq. B1)
MO2f = fish oxygen mass respiration (Eq. B2)
MOzp r = phytoplankton oxygen mass respira-
tion (Eq. 19)
MO2wcr = water column oxygen mass respira-
tion (Eq. 21)
MOzs r = sediment oxygen mass respiration (Eq.
22)
MOzd,z = effective oxygen diffusion at depth z
(Eq. B3)
MO2x s = excess 0 2 demand by sediment on
middle layer (Eq. 23)
N.B. All oxygen mass transfer terms in mg
O2/m3/hr.
The elements of the oxygen mass flux pre-
sented in Eq. 14 are shown diagrammatically in
Fig. 1. Equations for each of these mass flux
elements as implemented in the present model,
are shown in Appendix B. Revisions of previously
used formulations (Losordo, 1988) are described
below.
3.1. Phytoplankton oxygen production
In most ponds used for aquaculture (i.e. those
that are fertilized and can be considered eu-
trophic) phytoplankton provide the major source
and sink for dissolved oxygen. The productivity of
major phytoplankton species in the water column
are affected by many factors, including: intensity
of photosynthetically active solar radiation (PAR),
water column light attenuation, water tempera-
ture, p H and dissolved nutrient concentration.
These characteristics have themselves been the
focus of considerable study in limnology and
aquacultural science, and will not be discussed at
length here (see Losordo, 1980; Meyer, 1980;
Wetzel, 1983; Smith and Piedrahita, 1988; Boyd,
1990; Giovannini and Piedrahita, 1994).
In estimating the rate of phytoplankton oxygen
production, Steele's equation is commonly used
242 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
to describe this process (see Losordo, 1988 for
further treatment). Bannister (1974) presents
Steele's equation as follows:
MO2p = P * Chlorophyll - a concentration
( m g / m 3) (15)
P = (I/Imax)Pma x exp(1 - I/Ima×) (16)
where:
P = the rate of photosynthetic oxygen produc-
tion (mg O 2 / m g C h l - a / h r )
Pmax = maximum rate of oxygen production at
light saturation (mg O 2 / m g C h l - a / h r )
I = effective u n d e r w a t e r P A R intensity
(/.~mol/m2/hr)
Im~X= underwater P A R intensity at which light
saturation occurs (/.Lmol/m2/hr)
For use in the present model, Eq. 16 was
integrated over depth such that (Meyer, 1980):
Pvi = {[ Pmax exp(exp [ - ( I/Ima x) exp( - T/eZ 2)]
-exp[- ( I/Imax)exp(-~ezl)])]}
X {TIe(Z2 -- ZI)} -1 (17)
where:
Pvi = rate of photosynthetic oxygen production
for volume element ~ (mg O 2 / m g C h l - a / h r )
~Te = effective light extinction coefficient ( m - l )
z 1 = depth of the u p p e r boundary of volume
element V/ (m)
z 2 = depth of the lower boundary of volume
element V// (m)
The inclusion of this form of the primary pro-
ductivity equation for phytoplankton (Eq. 16) fol-
lows closely the approach used by Losordo (1988).
However, where Losordo conducted detailed pri-
mary productivity studies to determine the values
of Pmax and /max, the present model has used
several modifications in order to estimate these
values without resorting to extensive experimen-
tal studies.
In determining /max (the underwater P A R in-
tensity at which light saturation occurs), the pre-
sent model assumes that for locally adapted phy-
toplankton populations no light inhibition occurs
(Giovannini and Piedrahita, 1994). Therefore, the
value used for /max for a given simulation run was
set equal to the maximum P A R intensity recorded
at the pond surface at a particular site for a given
day. While this assumption may not be wholly
accurate, Losordo (1988) noted during his studies
that only rarely did the local solar regime exceed
his measured light saturation intensities, and even
then these intensities were found only in the very
top of the water column and for very short peri-
ods during maximum solar irradiance.
The inclusion of some sort of substitute for
Pmax is also needed for the model presented here,
since the P D / A CRSP research protocols do not
call for detailed primary productivity studies. The
value of Pmax, however, is integral to the correct
estimation of dissolved oxygen in aquaculture
ponds, and can be the source of much error in
D.O. prediction if it is estimated incorrectly. Epp-
ley (1972) presents an estimation for a term Ureax
which relates oxygen production to the rate of
carbon fixation by the phytoplankton. The value
of the carbon:chlorophyll-a ratio was initially
found from limits listed in Reynolds (1984), and a
calibration for the exact value of c a r b o n / c h l o r o -
phyll-a was conducted based on the known dis-
solved oxygen regime for a given pond. The val-
ues obtained for the ratio of carbon to chloro-
phyll-a fall well within the range of values re-
ported in Lee et al. (1991). The following relation
is used in the present model for estimation of
emax :
Pmax = ( C a r b o n / C h l o r o p h y l l - a) * Urea x (18)
where:
Carbon/Chlorophyll-a = 12.5-50 (from
Reynolds, 1984; calibrated for a given site and
day)
Umax = t e m p e r a t u r e - d e p e n d e n t maximum spe-
cific primary production rate (mg O 2 / m g C / h r ;
from Eppley, 1972)
3.2. Phytoplankton photorespiration
Respiration of the phytoplankton during pho-
tosynthesis has commonly been represented as
some proportion of the rate of photosynthesis
(Meyer, 1980; Losordo, 1988). While the present
model retains this somewhat crude assumption,
new evidence on phytoplankton respiration rates
provides a modification over the characterization
 S.D. Culberson, R. H. Piedrahita / Ecological Modelling 89 (1996) 231-258
offered previously. Giovannini (1994) reported
the presence of a time lag between the beginning
of the dark period (sundown) and the time at
which the phytoplankton respiration rate returns
to the baseline nighttime respiration rate. Based
on Giovannini's preliminary studies using high-
resolution, rapid-response oxygen probe studies
of phytoplankton populations, the time lag in the
present model is represented as three hours.
The rate of photorespiration is considered to
be 10% of the rate of photosynthetic oxygen
production (after Meyer, 1980 and Losordo,
1988):
MOzp r = 0.10(MOzv; D E L A Y 3.0 hr) (19)
where:
MO2p r = rate of D.O. respiration by phyto-
plankton (mg O 2 / m 3 / h r )
MOzp = rate of photosynthetic production by
phytoplankton (mg O 2 / m 3 / h r )
D E L A Y 3.0 hr = the time lag c o m p o n e n t
3.3. Water column respiration
The water column respiration rate ( W C R R ) is
defined in the current model as that rate of
respiration exhibited by the pond water column
not explicitly considered separately (Losordo,
1988). Included in this category are base respira-
tion of the phytoplankton (dark respiration), zoo-
plankton respiration, and respiration by sus-
pended bacteria. Estimations of this water col-
umn respiration rate are most conveniently made
using dark bottle incubations of water sampled
from the water column. For use in this model,
however, a slight revision of these estimations
was m a d e due to the fact that different water
column respiration rates could be calculated de-
pending on what time of day the samples were
collected, and on how long the dark bottles were
incubated (Teichert-Coddington and G r e e n ,
1992). In this revision, a regression equation was
generated from Teichert-Coddington and G r e e n ' s
data which related 4-hour, daytime-collected dark
bottle m e a s u r e m e n t s of water column respiration
rates to those rates measured by 4-hour, night-
time-collected dark bottle tests. In effect, the
daytime-collected m e a s u r e m e n t s overestimated
243
the water column respiration rates seen during
the nighttime hours. For the P D / A CRSP sites,
dark bottle estimations for nighttime water col-
umn respiration were calculated using darkbottle
m e a s u r e m e n t s obtained during the daytime, ad-
justed by the regression equation outlined above.
In addition, the water column respiration rate
was adjusted according to t e m p e r a t u r e (Krenkel
and Novotny, 1980) by:
kT1 = k T 2 * 0 (TI- T2) (20)
where:
k:rl = reaction rate at t e m p e r a t u r e T 1
k T 2 = known reaction rate at t e m p e r a t u r e T 2
0 = thermal coefficient (1.049; see Losordo,
1980)
As implemented in the present model water
column respiration is calculated as:
MOzwcr = MOzwcr m , 1.049 O'w~-to,) * 1000 (21)
where:
MO2wcr = W C R R of volume element, adjusted
for t e m p e r a t u r e (mg O 2 / m 3 / h r )
M O z w c r m = measured W C R R (mg O 2 / I / h r )
Tm = t e m p e r a t u r e at which W C R R was mea-
sured (°C)
There are, however, frequent cases where no
dark bottle m e a s u r e m e n t s of water column respi-
ration rates were available in the P D / A CRSP
Database. In these cases, water column respira-
tion rates are determined from the nighttime
decline in oxygen levels within the pond, and a
respiration term has been used which was derived
from a posteriori knowledge of the effect of this
respiration on dissolved oxygen levels in the pond.
The approach is less than ideal, but was useful in
testing the methodology and mechanistic accu-
racy of the revised model and not its absolute
predictive accuracy. What is hoped is that the
testing and validation of the model will reveal an
overall robustness and generality of the charac-
terizations of the physical, chemical, and biologi-
cal processes contained within the model, and
that the extension of these characterizations to
novel situations would have logical, accurate un-
derpinnings. Once the model has proven its abil-
ity to mimic current or historical aquaculture
244 S.D. Culberson, R.H. Piedrahita/ Ecological Modelling 89 (1996) 231-258
situations, it can be used to examine future or
" i m a g i n e d " situations.
3.4. Sediment respiration
Losordo's previous mechanistic water quality
model (Losordo, 1988) initializes the sediment
respiration rate c o m p o n e n t using measured sedi-
ment respiration rates, and subsequently adjusts
that rate using the t e m p e r a t u r e correction factor
as in Eq. 20. This rate was considered to be
constant over the entire pond bottom, and admit-
tedly was ill-defined. The P D / A CRSP research
sites do not routinely report sediment respiration
rates. Therefore, the nighttime dissolved oxygen
decline estimation for water column respiration
rates is considered to include that respiration
exhibited by the pond bottom sediments as well.
One slight modification to the estimation of
sediment respiration was made, however, in light
of recent work by fellow P D / A CRSP re-
searchers, and others (Madenjian, 1990; Szyper et
al., 1992). W h e r e sediment respiration rates were
measured and not included in the water column
respiration term, and where dissolved oxygen lev-
els fall below a certain threshold value (1.0 mg/1),
sediment respiration was assumed to decline to
zero. When this assumed threshold value was
omitted in the Losordo (1988) and present mod-
els, the lowest layer of the pond volume element
was frequently simulated as becoming anoxic,
even when the measured values in the pond for
this layer were 1.0 mg/1 or more.
As implemented in the present model sedi-
ment respiration is calculated as:
MO2s r = (MO2msr • O(Twc-Tm))/Yhs (22)
where:
MOzs r = sediment respiration rate, adjusted for
t e m p e r a t u r e (mg O 2 / m 3 / h r )
M O 2 m s r = measured sediment respiration rate
at measured t e m p e r a t u r e (mg O 2 / m Z / h r )
0 = thermal coefficient for benthic oxygen de-
m a n d (1.065; Krenkel and Novotny, 1980)
Th~ = thickness of sediment volume element
(m)
Following the example set by Losordo (1988),
the present model also accounts for excess sedi-
ment respiration resulting from the diffusion of
organic substances released into the overlying
water column by the benthic bacteria and other
processes contributing to oxygen demand. When
the D.O. declines to zero in the bottom water
volume element, the upward diffusion of these
organic substances may exert an additional oxy-
gen demand on the next higher volume element
(in the present model, the middle volume ele-
ment). This excess sediment respiration (in the
middle volume element) is calculated as (Losordo,
1988):
MO2x s = - (MO2p,bot + MO2d,z )
+ MOzs r * ( T h s / ( Z,b -- Zub)) (23)
where:
MOzx s = excess sediment oxygen demand (mg
O2/m3/hr)
MO2p,bot=photosynthetic oxygen production
for the bottom volume element (mg O 2 / m 3 / h r )
MO2d,~ = effective diffusion from middle vol-
ume element to bottom volume element (mg
O2/m3/hr)
Zub = depth of the upper boundary of the bot-
tom volume element (m)
Zlb = depth of the lower boundary of the bot-
tom volume element (m)
If the D.O. concentration of the bottom vol-
ume element is calculated to be greater than
zero, the value MO2× s is set to zero.
3.5. Dissolved oxygen model calculation procedure
In parallel with the calculation of model tem-
perature outputs described above, dissolved oxy-
gen calculations for each of the volume elements
involves tabulating the net oxygen change for
each element at each time step (Eq. 14). The net
oxygen change is then added to or subtracted
from the previous time step's oxygen mass, and
the current time step oxygen concentration is
solved for. Dissolved oxygen concentrations can
be calculated for any volume element (V/) at any
time (t) as:
DOvi a = DOvi,t_ , + ( d D O v i / d t ) * d t ) (24)
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258 245

where: tion for biologically-dependent factors is depen-

DOvi,t D.O. concentration of V~ at time t
= dent on the unique biochemical characteristics of
(mg/1)
DOvi,t_ 1 = D . O . concentration of V/ at time
t - 1 (mg/1)
d D O v i / d t = rate of change in D.O. concentra-
tion in V/ during the time interval ( m g / 1 / h r )
3.6. Input data summary and sample data set
The dissolved oxygen model uses as inputs
data collected during routine collection protocols
at P D / A CRSP research sites. A summary of the
data inputs needed for execution of the dissolved
oxygen portion of the model is given in Table 3.
All of the input values for the dissolved oxygen
c o m p o n e n t of the present model are those values
used for initialization. No time-series data be-
yond those used for the t e m p e r a t u r e simulations
are needed to produce predictions of dissolved
oxygen concentrations in the pond chosen for
simulation.
3. 7. Calibration for use at PD / A CRSP sites
Unlike the calibration method used in the de-
velopment of the t e m p e r a t u r e model, the calibra-
tion of the dissolved oxygen model was necessar-
ily pond specific - that is, while the solar radia-
tion and wind vector variables could be calibrated
using any pond on a given site (assuming these
factors are uniform over the whole site), calibra-
each individual pond. Typically, phytoplankton
populations show high variability in density from
pond to pond, and may also exhibit varying effi-
ciencies of primary production due to differences
in nutrient availability, light penetration, and
t e m p e r a t u r e stratification.
As mentioned, a calibration was needed for
the carbon : chlorophyll-a ratio. In the absence of
detailed primary productivity studies including
carbon:chlorophyll-a ratios like those conducted
by Losordo (1988), no acceptable substitute for
this calibration could be found. While this cali-
bration reduces the model's ability to make a
priori predictions of D.O. stratification in ponds
used for aquaculture, it does not lessen the
model's usefulness in examining the biochemical
mechanisms which determine dissolved oxygen
dynamics. The results obtained from the dis-
solved oxygen model following this pond-specific
calibration at each of the P D / A CRSP research
sites are presented below.
3.8. Simulation results from PD / A CRSP research
sites
3.8.1. Honduras
D a t a for the execution of the dissolved oxygen
model for the E1 Carao Aquaculture Experiment
Station were collected on Julian Day 351 (De-
cember 17), 1990, for Pond # B 7 (Table 3). The

Table 3
D.O. data inputs for CRSP P D / A research sites
Input variable Local time Honduras Rwanda Thailand
C a r b o n / C h l - a ratio 0600 45.45 28.57 166.67
Imax (tzmol/m2/s) N/A 1144 857.6 1543
Chl-a conc ( m g / m 3) 0600 234.6 33.0 20.8
Initial surface D.O. ( m g / l ) 0600 0.3 2.3 3.4
Initial middle D.O. (mg/1) 0600 0.2 2.3 3.4
Initial bottom D.O. (mg/1) 0600 0.2 2.3 3.4
Site elevation (m) N/A 583.0 1700.0 5.0
Total fish biomass (kg) 0600 26.68 12.9 12.0
Individual fish mass (g) 0600 200.0 63.0 43.0
W a t e r column resp rate ( m g / 1 / h r ) 0600 0.505 0.076 0.238
W a t e r column resp rate temp (°C) 0600 23.0 21.0 29.8
246 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258

Measured Dissolved Oxygen

Rwasave, Rwanda, collected on Julian Day 146
A 12 (May 26), 1989 for Pond #C1 (Table 3). The
resulting output for the three volume elements is
presented in Fig. 7, together with the measured
dissolved oxygen levels. As this figure shows, the
simulation for the Rwanda P D / A CRSP re-
search site is fairly accurate, especially for the
m
surface and middle water column volume ele-
ments. While there are some slight differences
O'e . . . . . . . . . . . between predicted and measured dissolved oxy-
0 4 8 12 16 20 24
gen levels, the character of D.O. dynamics is
captured quite well. Where there are differences
Simulated Dissolved Oxygen between simulated and measured values, they are
12 1.0 m g / l or less for the duration of the simula-
tion run. On the contrary, the character of the
bottom volume element is not at all well simu-
= 8 / / ~ • " Middle
lated, although the estimated D.O. concentra-
tions are within 1.25 m g / 1 of actual, and are
•0 4
usually closer than 0.75 mg/1. It is not entirely
clear to what factor this error in estimation can
0
o
m 2 be attributed for the bottom layer, but it is likely
O . . . . . . . . . . = = -- 1 . . . . .

0 4 8 12 16 20 24
Measured Dissolved Oxygen
Time (0 equals 0600 local hour)

Fig. 6. Measured and simulated D.O. profiles, Honduras,

Julian Day 351, 1990.

resulting simulation output for three depths is

shown together with measured dissolved oxygen
readings in Fig. 6. The surface D.O. levels are
predicted quite closely, but there is less accuracy
in the middle and bottom volume element esti- N 0
mations. The shape of the D.O. curve over time is 0 4 8 12 16 20 24
qualitatively correct in the middle element, but it
appears the phytoplankton primary productivity Simulated Dissolved Oxygen
is under-estimated within this element, since the 10
A
m a x i m u m predicted D.O. concentration is 2.0 ..I

mg/1 less than what was measured. The bottom £ 8 ----e--- SuflPace
Middle
volume element is poorly represented by the pre- • Bottom
6 '

sent model for this simulation, although it is clear

the model has indicated the presence of low D.O. o
"0
levels for all of the daylight hours. This indication > 1

is correct and useful from a practical pond man- 2'

m
agement standpoint. r~
- , - , - , . , .

4 8 12 16 2'0 24
Time (0 equals 0600 local hour)
3. 8. 2. R w a n d a
D a t a for the execution of the dissolved oxygen Fig. 7. Measured and simulated D.O. profiles, Rwanda, Julian
model for the Rwasave P D / A CRSP site at Day 146, 1989.
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258 247

Measured Dissolved Oxygen

the phytoplankton primary productivity charac-
15'
terization is again at fault. T h e r e may also be
some influence of altitude (the Rwasave Station E 12"
is located at 1700 meters elevation) on the pro- =.
cesses within the pond which the present model 9'

does not take into account.

6'

3.8.3. Thailand | 3.
D a t a for the execution of the dissolved oxygen 5
c o m p o n e n t of the present model for the Ayut- 0 . , - , -

0 4 8 12 16 2'0 24
thaya Station facility in Thailand were collected
on Julian Day 96 (April 6), 1988, for Pond # 1 2
(Table 3). Predicted dissolved oxygen outputs and Simulated Dissolved Oxygen
the corresponding measured D.O. levels for this
date and pond are shown in Fig. 8. The qualita-
tive and quantitative accuracy of this simulation is
-'5 t Sulr/'ace
= 1 ~ • Mk~
quite high. The present model is only slightly
errant when estimating maximum surface D.O.
concentration (approximately 1.5 m g / 1 too low).
The character of the dissolved oxygen dynamics
in all three volume elements is captured well, and
the hourly predictions of D.O. concentration are
0 4 8 12 16 20 24
very close to the measured values. The accuracy Time (0 equals 0600 local hour)
of the Thailand simulation run may be related to
Fig. 8. Measured and simulated D.O. profiles, Thailand, Ju-
the overall completeness of the Ayutthaya Station
lian Day 96, 1988.
portion of the P D / A CRSP Database.

3.9. Comparison of actual and simulated D.O.
concentration profiles
Table 4 presents a summary of the actual
timing and extent of dissolved oxygen stratifica-
tion in the ponds described above, and the corre-
sponding model prediction of these same events.
As can be seen from the table, and in common
with the t e m p e r a t u r e simulations, the qualitative
nature of the simulations is rather accurate; the

Table 4
Comparison of actual and predicted D.O. stratification events
Site and Julian Stratification Maximum Isothermal Maximum
pond day onset a stratification conditions b stratification c
Actual Simulated Actual Simulated Actual Simulated Actual Simulated
(h) (h) (h) (h) (h) (h) (mg/l) (rag/l)
Honduras
Pond#B7 351-352 0800 0900 1430 1500 2200 1900 11.2 9.7
Rwanda
Pond #C1 146-147 0900 1100 1600 1600 0200 0500 1.3 3.0
Thailand
Pond #12 96-97 0800 0900 1600 1500 2200 2400 7.3 7.1

a D.O. differences greater than 1.0 m g / l are considered stratified.

b D.O. differences less than or equal to 1.0 m g / I are considered isotropic.
c Measured as the difference between surface and bottom readings.
248 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
onset of stratification, the timing of maximum
stratification, and the resumption of fully-mixed
conditions within the ponds at all three sites are
closely predicted. Less accurate were the predic-
tions of individual dissolved oxygen levels in the
volume elements at each hourly interval, although
for the maximum stratification events, the model
was within 1.13 m g / l of oxygen, averaged over
the three ponds.
3.10. Discussion: Dissolved oxygen
The predictive accuracy of the dissolved oxy-
gen model remains quite high in spite of the
mass-balance modifications and model simplifica-
tions. While there are some slight differences
between the measured and modelled dissolved
oxygen concentrations from the P D / A CRSP ex-
perimental sites in terms of absolute values, the
qualitative nature of the model outputs mimic
quite well those dynamics encountered within the
ponds - the timing of oxygen peaks, the magni-
tude of the peaks, the extent of oxygen stratifica-
tion, and the simulation of anoxic conditions near
the bottom of the pond are all represented quite
accurately.
The present model, initialized and calibrated
with fairly simple and easily-obtainable weather
and water quality data, can be a reliable tool to
be used when trying to understand if changes in
pond management or morphology will have quali-
tative a n d / o r quantitative effects on dissolved
oxygen levels within the pond. Also important to
note is the robustness of the model when simulat-
ing ponds located in different parts of the world:
the outputs presented here range from coastal to
inland tropical regions, and from elevations near
sea level to the mountains of East Africa (eleva-
tion 1700 meters).
4. Simulation experiments
Using the Aquaculture Pond Ecosystem Model
(A.P.E.M.) described above, a series of simula-
tion runs were conducted to investigate the ef-
fects of different management strategies on pond
temperature and dissolved oxygen regimes. The
baseline run used was that for the Thailand P D / A
CRSP site described previously (Tables 1 and 3).
Pond depth and secchi disk depth (S.D.D.) for
the baseline run were 0.96 m and 0.35 m, respec-
tively. The model was used to examine the effect
on dissolved oxygen and temperature stratifica-
tion of changes in pond depth and secchi disk
depth. The model was run for pond depths of
0.48 m, 0.96 m, and 1.92 m, and for secchi disk
depth to pond depth ratios of 0.25, 0.50, and 1.0.
Secchi disk depth measurements are frequently
used to estimate phytoplankton density in aqua-
culture ponds (at least on a relative basis), and
changes in S.D.D. represent changes in phyto-
plankton concentration. Since phytoplankton
density is assumed to be the most important
factor influencing pond D.O. concentrations
(which implicitly incorporates management prac-
tices such as fertilizer application rates, water
exchange frequencies, cropping effects associated
with fish feeding, etc.), the results generated by
the A.P.E.M. may therefore be thought of as
indicative of overall pond behavior, and can be
assumed to apply more generally to ponds other
than the baseline pond selected for illustration
here. Details of the simulation runs executed are
examined below.
4.1. Secchi disk depth/phytoplankton density ra-
tios
Changes in secchi disk depth are assumed to
correlate with changes in the density of the phy-
toplankton standing crop. In order to develop
reasonable estimates of phytoplankton crop den-
sity for each of the hypothetical S.D.D. inputs, a
simple regression was made for the pond under
scrutiny using measured values of S.D.D. and
phytoplankton density taken on other dates from
this pond ( P D / A CRSP, 1992; Fig. 9). The re-
gression only holds for S.D.D. measurements
shallower than approximately 0.44 m. Where the
hypothetical S.D.D. measurements used as inputs
to the model exceed 0.44 m, the phytoplankton
densities (expressed as concentration of chloro-
phyll-a) are reduced to a default value of 5.0
m g / m 3. The use of a default value of 5.0 m g / m 3
chlorophyll-a for all S.D.D.s greater than 0.44 m
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258 249

A temperatures but tend to have cooler tempera-

25"
CZD .3910x R^2 = 0.808 tures in deeper waters.
g 20" In general, the shallower ponds are less strati-
,= fied than the deeper ponds (Fig. 10 versus Figs.
15'
11 and 12), and the more turbid ponds are more
D.
10'
o stratified than clearer ones (Figs. 10a, l l a , and
o
x.- 5' 12a). This is in agreement with what is expected,
o
0
given the light extinction coefficients associated
25 30 35 40 4'5 with the various turbidities (expressed as S.D.D.).
Secchi Disk Depth (cm)
The pond represented in Fig. 12a (pond depth
Fig. 9. Chlorophyll-a/Secchi disk depth relation, CRSP Pond 1.92 m; S.D.D. 0.48 m) shows the pond remaining
#C12, Thailand. stratified over the 24-hour simulation period, a
p h e n o m e n o n associated with larger, deeper bod-
ies of waters such as lakes (Goldman and Horne,
in effect implies that as S.D.D. increases beyond 1983) - this corresponds to what one might ex-
0.44 m, non-algal turbidity decreases. The deeper pect given a deeper water column and a relatively
S.D.D. values used (0.48 m, 0.96 m, and 1.92 m) shallow S.D.D.
are therefore associated with lower concentra-
tions of dissolved and non-algal particulate sub-
stances that may absorb light.
a: SDD 0.12m
4.2. Simulation results: Temperature
32
The temperature results from the simulations
31
discussed are shown in Figs. 10-12. The most
3O
thermally-homogeneous pond generated through
29
the simulations is that with a shallow depth (0.48
28 .
m) and a correspondingly deep S.D.D. (0.48 m) 0 4 8 12 16 20 24
(Fig. 10c). It is unlikely, however, that this pond b: SDD 0.24m
would be useful in aquaculture, since the low 34
phytoplankton density would also reflect low food o
Surface
v

availability for herbivorous culture animals. An- 32

other difficulty with such a management scheme = 31
(0.48 m pond depth; 0.48 m S.D.D.), is the likeli- 20.
hood of rooted macrophytes invading the pond, E 29
k-
which ultimately may prove costly to eradicate or 28
control. Nearly as homogeneous as this pond is
the pond depicted in Fig. 10b, where pond depth c: SDD 0.48m
is 0.48 m and S.D.D. is 0.24 m, a simulation
which combines a relatively shallow pond with a
moderate phytoplankton bloom. This shallow
31
depth and moderate phytoplankton density seems
3O
to provide sufficient conditions for minimizing
29 _ ~. _
temperature stratification, while at the same time
28 ,
maximizing overall primary productivity. A draw- 0 4 8 12 16 20 24
Time (0 e a u a l s 0 6 0 0 local h o u r l
back of this pond depth and S.D.D. combination
is the uniformly high temperature reached in the Fig. 10. Temperature simulation results for pond depth of
pond. Deeper ponds may reach similar surface 0.48 m.
250 S.D. Culberson, R.H. Piedrahita /Ecological Modelling 89 (1996) 231-258
a: SDD 0.24m
mately 5 mg/1 in Fig. 13a and 6 m g / l in Fig.
33 13b). Conversely, for the lowest phytoplankton
32 density (Fig. 13c), there is negative net oxygen
production within the pond. In this last case
there does not seem to be enough phytoplankton
in the pond to provide for the oxygen needs
included in the various respiration terms.
0 4 8 12 16 20 24
For the pond of simulated depth equaling 0.96
b: SDD 0.48m m and S.D.D. of 0.24 m (Fig. 14a), the dissolved
oxygen profiles are similar to those of the base-
o 33 "'-9-- Surface
• M dle line run. In general, this configuration (0.96 m
• B o t t o m
pond depth and 0.24 m S.D.D.) results in the
most stratified D.O. levels - up to a 7 or 8 m g / l
D,,
difference between the surface and bottom lay-
E
~ . , . , , . , . , .
ers. This configuration could have considerable
0 4 8 12 16 20 2'4 effect on the ability to maximize production of
bottom-dwelling species like shrimp. For the
c: SDD 0.96m
less-dense phytoplankton densities in this pond

31 a: SDD 0.48m
3O

0 4 8 12 16 20 24
Time (0 equals 0600 local hour)

Fig. 11. T e m p e r a t u r e simulation results for pond depth of

0.96 m.
~ -- . . . . . . . . . . .

0 4 8 12 16 20 24

b: SDD 0.96m

---e-- Surface
4.3. Simulation results: Dissolued oxygen s Middle
• 32 • Bottom
The D.O. results from the simulations outlined = 31'
above are shown in Figs. 13-15. For the pond
with the simulated depth of 0.48 m, the interme-
1-
diate situation (Fig. 13b), where S.D.D. is 0.24 m, 28
4 8 12 16 2'0 24
yields the most satisfactory results, both in terms
of limiting D.O. stratification during the middle c: SDD 1.92m
of the simulated day and in maximizing total 34
33
primary productivity for the simulated 24-hour
32
period. The most-dense phytoplankton case for
31
this pond (Fig. 13a) shows some detrimental ef-
30
fects caused by net phytoplankton respiration in
2 9 - - -
the bottom layer of the pond (since it is likely the
2 8 , " " " '
phytoplankton in this layer are light-limited), 0 ; 8 12 1'6 2'0 24
Time (0 eauals 0600 local hour)
which also has the effect of lowering the net
oxygen production of the pond (represented by Fig. 12. T e m p e r a t u r e simulation results for pond depth of
the D.O. after the 24-hour simulation: approxi- 1.92 m.
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258 251

a: SDD 0.12m
15- depth increases. Similarly, as the turbidity of the
water increases (signified by lower S.D.D), so
12-
does the tendency for stratification. In the analy-
9-
sis presented here, a low S.D.D. is associated
6- with high concentrations of phytoplankton. These
3; phytoplankton are net oxygen producers when
they receive adequate amounts of light (close to
o " ; " ; ;2 1; 20 2',
the surface of the pond), but can become net
A b: SDD 0.24m oxygen consumers in deeper areas of the pond
15" where oxygen production is insufficient to meet
12" the respiratory needs of the plankton.
~ 9" Of the pond configurations tested, a pond with
X

0 6"
'
a depth of 0.48 m and a S.D.D. of 0.24 m resulted
in the highest net oxygen production and the
• 3
least temperature and D.O. stratification. This
0 4 8 12 1 '6 2'0 24 configuration contrasts with the measured values
for the Thailand pond used as a reference for the
c: SDD 0.48m
simulations (0.96 m depth and 0.35 m S.D.D.).
One characteristic of particular note, however, is
12

a: SDD 0.24m

0
0 4 8 12 16 20 24
'!t
1

Time (0 eauals 0600 local hour)

Fig. 13. Dissolved oxygen simulation results for pond depth of

0.48 m.
0 4 8 12 16 20 24

A b: SDD 0.48m
(Fig. 14b and c), some stratification still is seen
during the middle of the day, but the phytoplank-
12 ] • Middle
ton densities simulated would not be adequate to
meet the oxygen demands of intensive aquacul-
ture.
Similar results are obtained for the final pond
depth simulated (1.92 m; Fig. 15a, b, and c). For O I - , - , - , - , - , -

0 4 8 12 16 20 2'4
the 1.92-m-deep pond simulations, only in the
final case (S.D.D. of 1.92 m) does the pond c: SDD 0.96m
remain aerobic in the bottom layer.

4.4. D&cussion." Simulation experiments

Results obtained from the simulations pre-

sented here represent the effects of pond depth
,

and turbidity changes on temperature and dis- 0 4 8 12 16 20 24

solved oxygen regimes in aquaculture ponds. As Time (0 equals 0600 local hour)

can be seen in Figs. 10 through 15, the tendency Fig. 14. Dissolved oxygen simulation results for pond depth of
and stability of stratification increases as pond 0.96 m.
252 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258

a: SDD 0.48m
increases in depth result in lower temperatures in
the surface layer, but in more stable stratification.
1
For example, where temperature uniformity for
the 0.48-m-deep pond with a S.D.D. of 0.12 m
(Fig. 10a) was reached at approximately 1700
3 hours local time, the equivalent but deeper pond
in Fig. 12a (1.92 m deep, 0.48 m S.D.D.) re-
0 4 8 12 16 20 24 mained stratified until 0500 hours the next day,
b: SDD 0.96m the end of the simulation period.
The amount and distribution of oxygen pro-
12 - duced in the water column depends on light
intensity and penetration, as well as on phyto-
plankton concentration. The empirical relation-
ship between light penetration (S.D.D.) and phy-

0 t-
0 4 8 12 16
C: SDD 1.92m
15'
12'
9"
6"
0
o " ; " 8 " ,'2
Time (0 equals 0600 local houri
Fig. 15. Dissolved oxygen simulation results for pond depth of
1.92 m.
the large temperature fluctuation within the pond
over the 24-hour simulation cycle (Fig. 10b). This
pattern of large temperature fluctuations is ob-
served in all the shallow ponds when compared to
deeper ponds having similar S.D.D. values. These
larger temperature fluctuations are due to the
smaller total thermal mass of water in the shal-
lower ponds. Similarly, a low S.D.D. value for a
given pond depth results in more marked overall
stratification and higher temperature fluctuations
in the surface layer of the pond. In general, as
pond depth and S.D.D. increase, so does the
overall uniformity in water temperature. Con-
versely, for a given S.D.D., deeper ponds tend to
stratify more than shallower ponds.
For ponds with equal ratios between pond
depth and S.D.D. (e.g. Figs. 10a, 11a, and 12a),
20
,
24
toplankton concentration used in the simulations
presented here is shown in Fig. 9. A residual
phytoplankton concentration of 5 m g / m 3 (as
chlorophyll-a) was assumed for S.D.D. values
greater than 0.44 m as explained above. As a
result of this assumption, ponds simulated with a
S.D.D. of 0.48, 0.96, and 1.92 m were considered
to have a phytoplankton chlorophyll-a concentra-
tion of 5 m g / m 3. Dissolved oxygen concentration
and stratification in the ponds varied with both
2'0 2', pond depth and S.D.D. (Figs. 13-15). Dissolved
oxygen in the bottom layer, and overall oxygen
fluctuation over the diel cycle were highest in the
0.48-m pond. By contrast, D.O. in the bottom
layer reached zero for two of the S.D.D. tested
with the 1.92-m-deep pond (Fig. 15a and b). In
the shallow pond, an increase in S.D.D. from 0.12
to 0.24 m resulted in a decrease in stratification,
and increases in both the peak D.O. concentra-
tion and overall oxygen production for the 24-hour
cycle (Fig. 13a and b). Further increasing the
S.D.D. resulted in a drop in phytoplankton con-
centration to 5 m g / m 3 chlorophyll-a, and an
overall drop in D.O. production. Dissolved oxy-
gen concentration in the 0.48-m-deep pond with a
S.D.D. of 0.48 m was therefore lower than for the
other two S.D.D. values used (Fig. 13a, b and c).
D.O. stratification was most marked in the
0.96-m pond with a S.D.D. of 0.24 m (Fig. 14a).
The dissolved oxygen difference between surface
and bottom layers reached approximately 8 mg/1,
and remixing of the layers occurred late in the
day for the surface and middle layers (1800 hours
local time), and in the early morning for the
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
bottom layer (about 0200 hours). Dissolved oxy-
gen concentration in the bottom layer of the
0.24-m-S.D.D. pond (Fig. 14a) reached its lowest
value shortly after midnight, and just prior to
mixing with the layers above it. The timing of the
low D.O. values in the bottom layer is particularly
important for shrimp and other benthic animal
culture systems.
Dissolved oxygen concentrations in the 0.96-m
pond was lowest for 0.48 m S.D.D., and increased
slightly when S.D.D. increased to 0.96 m (Fig. 14b
and c). This rise is caused by the increase in
oxygen production caused by the improved light
penetration with the same phytoplankton concen-
tration of 5 m g / m 3 chlorophyll-a, and with the
same sediment and fish oxygen demand in both
cases.
Dissolved oxygen production in the 1.92-m
pond was low for all S.D.D. values tested due to
the low phytoplankton concentration (Fig. 15a, b
and c). The bottom layer in the ponds with S.D.D.
values of 0.48 and 0.96 m did not mix with the
middle layer, and became anaerobic sometime
during the night (Fig. 15a and b). By increasing
the S.D.D. in this pond to 1.92 m, an increase in
oxygen concentration in the pond was obtained
due to the rise in light penetration and the drop
in non-algal turbidity while maintaining constant
phytoplankton chlorophyll-a concentration. While
a moderate degree of stratification was observed
in the 1.92-m-deep, 1.92-m-S.D.D. pond, dis-
solved oxygen concentration remained low
throughout the diel cycle, never reaching above 6
mg/l, and dropping to just under 2 m g / l at the
end of the 24-hour simulation.
The low D.O. values in the shallow pond (0.48
m) with a high S.D.D. (0.48 m; Fig. 13c), when
compared to other ponds with S.D.D. equal to
their pond depths (Fig. 14c and Fig. 15c), was the
result of the low overall oxygen production rela-
tive to the demand by the sediments and fish
crop, which were the same for all the ponds
tested. By increasing the depth of the pond and
of the S.D.D., and maintaining phytoplankton
chlorophyll-a concentration at 5 m g / m 3, the
overall oxygen production was increased. This
increase resulted in a rise in the D.O. available
and in the D.O. concentrations in the ponds.
253
5. Conclusions
The results from this Aquaculture Pond
Ecosystem Model should not be applied unre-
servedly across all aquaculture sites. This model
and these simulations are an attempt to quantify
the chemical, physical, and biological processes
occurring within an aquaculture pond, and then,
having achieved an approximation of the pond
ecosystem (the working model), make certain rea-
sonable changes in inputs and functions to inves-
tigate the probable effects these changes may
have on the culture ecosystem. Model outputs
illustrate the complexity of responses that result
from simple changes in pond management pa-
rameters; depth and S.D.D. in this case. The
situations presented represent a particular com-
puter "experiment". Similar experiments can be
conducted for other situations prior to carrying
out field trials. The speed, ease, and inexpensive
nature of using a model should appeal to anyone
interested in making better use of resources when
striving for improved aquaculture methods and
management techniques. This model does not
presume to replace the careful experimentation
performed every day by practicing aquaculturists,
but does provide a complement to these efforts.
Making models and asking hypothetical questions
(which should, after all, arise during the culture
cycle anyway) is one way to reduce the cost and
time involved in refining our management tech-
niques. The use of pond ecosystem models could
drastically reduce the need for lengthy experi-
mental trial culture cycles by more closely focus-
ing on what the optimum culture conditions for a
given site might be, as suggested by simulation
results.
The simplifications made within the present
model and the increased user-friendliness with
which it is constructed make available a basic
pond ecosystem model which can incorporate
more accurate characterizations of phytoplankton
physiology and pond ecosystem behavior, as these
characterizations become available. This basic
temperature and dissolved oxygen model also
provides a framework into which may be added
characterizations of pH, alkalinity, nutrient dy-
namics, and ultimately fish yield.
254 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
Acknowledgements
The research described herein has been con-
ducted through funding by the United States
Agency for International Development, under
grant DAN-4023-G-00-0031-00. The P D / A CRSP
accession number for this paper is 133.
The efforts of all P D / A CRSP personnel in-
volved in the conduct of experiments and the
collection of water quality and related data are
specially acknowledged.
Additionally, the authors would like to thank
Dr. Tom Rumsey, Professor of Biological and
Agricultural Engineering, University of California
at Davis, for suggestions and comments regarding
the estimation of daily air temperature profiles.
Earlier versions of parts of this paper were
presented at the 1992 Annual Meeting of the
World Aquaculture Society, Orlando, in a session
co-sponsored by the American Society of Agricul-
tural Engineers (ASAE Paper No. A Q U A 92-
102), and at Techniques for Modern Aquacul-
ture, a conference sponsored by the American
Society of Agricultural Engineers in Spokane,
Washington, June, 1993.
Appendix A. Temperature model equations
Equations as used by Losordo and Piedrahita
(1991) and as included in the present model. The
original source of the expression is indicated.
(~sn = ~fls -- ~)r "~- Cs(1 -R)
(WRE, 1968; Losordo and Piedrahita, 1991),
0r = 6 ~ ( R ) ;
~b~ = the measured incident radiation ( k J / m 2 / h r ) ;
R = Rs(1 - (0.08Wz))
(Losordo and Piedrahita, 1991);R=reflectivity
adjusted for surface roughness; Wz = wind veloc-
ity at height z meters above pond surface (m/s);
R s = 2.2(180A/~-)-0.97
(WRE, 1968); A = solar altitude angle.
6at = (1 - r)etr(Tak) 4
(Octavio et al., 1977), where:r = r e f l e c t a n c e of
the water surface to longwave radiation --0.03
(decimal fraction); o-= Stefan-Boltzman con-
stant, 2.07 × 10 -7 ( k J / m Z / h r / K 4 ) ; Tak = absolute
air temperature above the water surface (K);
e = emittance of the atmosphere
= 0.398 X 10-5(Zak) 2"148 (dimensionless)
(A2.1)
(Octavio et al., 1977).
&ws = 0.97o,(Twk) 4 (A3)
(Octavio et al., 1977; Henderson-Sellers, 1984),
where: Twk = temperature of the water in K.
4)c = 1.5701W2( Two- Tac) (A4)
(Fritz et al., 1980), where: W2 = wind speed at a
height of 2 m above the pond (m/s); Twc=
temperature of the water at the surface (°C);
Tac = temperature of the air at the surface (°C).
q~sn,z = qbsn(1 - R ) ( 1 - / 3 ) exp( - r / e Z ) (A5)
(Laska, 1981; Stefan et al., 1983; Lund and
Keinonen, 1984; Cathcart, 1987; Losordo and
Piedrahita, 1991), where: ~bs,,z = solar irradiance
at depth z (kJ/mZ/hr); /3 = fraction of incident
solar irradiance absorbed at surface = 0.03 (frac-
tion); r/e = effective light extinction coefficient
(m- 1);
~/e = r / = 1 . 7 / S D D (A5.1)
(A1) (Poole and Atkins, 1929) where SDD = secchi
disk depth (m).
(AI.1) q~sed = ksed((Tbot- Tsea)/Az) (A6)
(Losordo and Piedrahita, 1991), where: kse d =
thermal conductivity coefficient for sediment--
(A1.2) 2.53 (kJ m -1 h -1 °C -1) (Hull et al., 1984); Tbot =
temperature of the bottom water volume element
(°C); Tsed = temperature of the sediment volume
element (°C); Az = distance between the centers
of the volume elements (m).
(A1.3)
~bgw= ke((Tsed - Tgw)/Az ) (A7)
(Losordo and Piedrahita, 1991), where: k ~ =
(m2) thermal conductivity coefficient for the earth =
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
2.53 (kJ m -1 hr -1 °C -1) (Hull et al., 1984);
Tgw = t e m p e r a t u r e of the ground water = 20.0
(°C); Az = distance between sediment and ground
water volume elements = 5.0 (m).
q~d,z=pw. Cpw* A v * E z , z ( A T / A z )
(after Chapra and Reckhow, 1983), where:pw =
density of water (kg/m3); Cpw = heat capacity of
water = 4.1816 (kJ K -1 kg-1); A v = idealized
volume element surface area = 1.0 (m2); Ez, z =
effective diffusion coefficient at depth z; A T / A z
= t e m p e r a t u r e versus depth gradient between
volume centers of adjacent volume elements;
pw = (0.99987 + ( ( 0 . 6 9 × 10-5)(Twc))
- ((8.89 × 10-6)(Twc) 2)
+ ( ( 7 . 4 × 10-8)(Twc)3))* 1000 ( k g / m 3)
(Spain, 1982);
Ez, z = ( Eo,z)(1 + 0.05Riz) -1
(Sundaram and Rehm, 1973);
Eo, z = [(Ws * )2//zsk *] exp( - k *z)
(Henderson-Sellers, 1984), where:
W s * = frictional velocity due to wind stress
(m/s);
/z~ = drift velocity ( m / s ) ;
k * = decay coefficient (m-1);
WS* = ( To/Pw ) 1/2
(Henderson-Sellers, 1984);
'ro=Pafz(WVz) 2 (N/m 2)
(Henderson-Sellers, 1984), where:
Pa-----density of air above the water surface =
1.1988 (kg/m3);
C z = coefficient of aerodynamic resistance =
1.0 X 10 -3 (Losordo, 1988);
Wv z = wind vector magnitude at a height of z
meters above the pond ( m / s ) ;
/x s = 3 0 ( W s * ) (m/s)
(Henderson-Sellers, 1984);
k* = 6W~ 1.84
255
(Henderson-Sellers, 1984), where Wsa = wind
shear area = wind speed × length of wind fetch
(see Smith, 1979);
Ri~ = [ a v * g ( z ) Z / ( W s * ) Z ] ( A T / A z ) (A8.8)
(A8)
(Sundaram and Rehm, 1973);
a v = 1.5 × 10-5(Tav - 277)
- 2.0 X 10-7(Tav - 277) 2 (A8.9)
(Sundaram and Rehm, 1973), where:
Ez, z = the effective diffusion coefficient at
depth z, adjusted for water column density
gradient (mZ/hr);
Ri z = Richardson n u m b e r at depth z (dimen-
sionless);
a v = coefficient of expansion for water (K-1);
g = gravitational acceleration constant = 9.81
(A8.1)
(m/s/s);
z = depth for which the Richardson n u m b e r is
being calculated;
AT = t e m p e r a t u r e difference between adjacent
(A8.2) volume elements (K);
Az = distance between adjacent volume ele-
ment centers (m);
(A8.3) Tar = average water t e m p e r a t u r e of the adja-
cent volume elements (K).
Appendix B. Dissolved oxygen model equations
Equations as used by Losordo (1988) and in-
(A8.4) cluded in the present model. The original source
of the expression is as indicated.
(A8.5) B.1. Oxygen diffusion t o / f r o m the atmosphere
Diffusion of oxygen to or from the water sur-
face and the overlying atmosphere depends upon
the difference between the saturation concentra-
tion of oxygen in the pond water and the actual
dissolved oxygen concentration present in the
surface water layer. The mass rate of oxygen
transfer has been shown to be a function of an
(A8.6)
oxygen transfer coefficient (Tchobanoglous and
Schroeder, 1985) as:
(AS.7) MO2D = K ( C s - C) * 1 0 0 0 . 1 / T v e (B1)
256 S.D. Culberson, R.H. Piedrahita/ Ecological Modelling 89 (1996) 231-258

where: M O 2 D = the rate of diffusion across
a i r / w a t e r interface (mg O 2 / m 3 / h r ) ; K = oxygen
transfer coefficient ( m / h r ) ; C s = saturation con-
centration of D.O. in water at a given elevation
and t e m p e r a t u r e (mg O 2 / 1 ) ; C = concentration of
D.O. present in water (rag O 2 / 1 ) ; Tve = thickness
of the surface volume element (m); and
K = 0.0036(8.43 * (Wz) °5 - 3.67 * (Wz)
+ 0 . 4 3 . (Wz) 2) (BI.I)
(Banks and Herrera, 1977), where: Wz = w i n d
speed at reference height z = 2 m ( m / s ) .
Csr = 14.625 - 0.41022. (Twc)
+ 0.007991 *(Twc) 2
- 0.0000778 * (Two) 3 (B1.2)
(Krenkel and Novotny, 1980), where: Csv =
saturation concentration of D.O. in water at tem-
perature Tw¢ (mg O2/1); Twc = water t e m p e r a t u r e
(°C).
Cs = Csr * (1 - 0.0001 * E ) (B1.3)
(Krenkel and Novotny, 1980), with: E = site ele-
vation (m).
B.2. Fish respiration
MO2f = F r * F u (B2)
(Losordo, 1988), where:
Fr = (10 (x)) * 1000 (B2.1)
(Boyd, 1990), where:
x = - 0.999 - 9.57 * 10 - 4 . wt + 6.0 * 10 - 7 . (wt) 2
+ 3 . 2 7 . 1 0 -2 * Two - 8 . 7 . 1 0 -6 * (Twc) 2
+ 3.0 * 10 - 7 *wt* Two
Fr = respiration rate per weight of fish (mg
O 2 / k g / h r ) ; wt = average fish weight (g).
F b = FW/(A * d) (B2.2)
where: F b = average fish biomass concentration
(kg/m3); FW = total fish biomass (kg); A = pond
surface area (m2); d = depth (m).
Inputs of fish biomass are based on periodic
sampling conducted at the P D / A CRSP research
sites. In addition, average weights of fish are
classified into three groups, and the decimal frac-
tion of these groups (termed 'small,' 'medium,'
and 'large') is estimated by field personnel (al-
though for the simulations presented here all of
the fish were of approximately uniform size, and
were all lumped into the small category). Respi-
ration rates are calculated independently for each
of the fish size classes, and the total fish respira-
tion term is arrived at by summing the three size
class terms (Losordo, 1988).
In order to account for the fact that fish will
avoid areas of low dissolved oxygen in ponds
(Gebhart and Summerfelt, 1976) the present
model also retains the Losordo (1988) method of
simulating vertical migration by the fish away
from volume elements which have low D.O. con-
centrations. At each time step the calculated dis-
solved oxygen concentration is compared with an
input term " L O W D O " , beginning with the bot-
tom volume element. If the calculated D.O. con-
centration is below L O W D O , the model shifts
the biomass of the fish in the bottom element to
the middle volume element. The same procedure
is then repeated for the middle volume element.
In the event that the calculated dissolved oxygen
concentration in all three volume elements is
below L O W D O , all of the fish biomass is consid-
ered to reside in the surface volume element.
B.3. Oxygen diffusion within the water column
Effective diffusion of oxygen in the present
model is assumed to follow that of energy diffu-
sion described for the t e m p e r a t u r e model. While
there might be other ways in which to conceptual-
ize and define the effective diffusion of oxygen
within the pond water volume, the successful use
of this assumption by Losordo (1988) recom-
m e n d e d identical treatment of oxygen diffusion
in the present model. The calculation of the
diffusion of oxygen between water volume ele-
ments is therefore:
MO2d,z = Ez, z * ( D O 1 - D O 2 )
/ ( zc2 - zcx) * 1000 * 1 / t h (B3)
 S.D. Culberson, R.H. Piedrahita / Ecological Modelling 89 (1996) 231-258
where:
Ez z = effective diffusion coefficient at depth z
(m2/hr);
DO1 = D.O. concentration in volume element
1 (mg/1);
DO 2 = D.O. concentration in volume element
2 (mg/1);
Z¢l = d e p t h o f t h e c e n t e r o f v o l u m e e l e m e n t 1
(m);
zc2 = d e p t h o f t h e c e n t e r o f v o l u m e e l e m e n t
(m);
th = thickness of the volume element into which
o x y g e n d i f f u s e s (m).
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