Aquacultural Engineering 1 (1982) 245-261

COMPUTER MODELING OF THE DIURNAL OXYGEN LEVELS

IN A STILLWATER AQUACULTURE POND

D. I. MEYER
Department of Electrical Engineering, University of Calijbrnia, Davis, CA 95616, USA
and

D. E. BRUNE
Department of Agricultural Engineering, The Pennsylvania State University, University Park,
PA 16802, USA

ABSTRACT

A dynamic model o f dissolved oxygen behavior in a stillwater aquaculture pond is

presented. Using theories and principles that have been established for aerobic waste-
water treatment ponds and shallow lakes and reservoirs, equations were developed to
describe the short-term dissolved oxygen fluctuations o f an aquaculture pond. Com-
ponents considered in this model include the consumption and production o f oxygen
by phytoplankton, fish, detritus, and the process o f reaeration. Factors affecting these
components can be specified by the user; these blclude meteorologic and geographic
data, and phytoplankton, .fish, and detrital characteristics. After comparing the model
to field data and analyzing the sensitivity o f its components, areas o f research which
may have an impact on pond management are identified.

NOMENCLATURE

AP average photosynthesis rate

AR surface area of pond (m 2)
A TM atmospheric attenuation factor (dimensionless)
BODT BOD remaining after one day period
BODL total BOD at beginning of decay period
CL cloudiness factor (dimensionless)
D water depth (m)
245
Aquaeultural Engineering 0144-8609/82/0001-0245]$02.75 © Applied Science Publishers Ltd,
England, 1982
Printed in Great Britain
246 D. I. MEYER, D. E. BRUNE

D1, De boundaries of water column (m)

DEC declination of the sun (radians)
DT time of day (h)
EC light extinction coefficient (ly/m)
ELIT effective light intensity (ly)
ER endogeneous respiration rate
EXT light extinction coefficient (m -1)
FR fish oxygen demand as function of OP
FRO fish oxygen demand as function of FW and WTMP
FW fish weight
1SL initial slope of photosynthesis versus light intensity
JD Julian date (days)
KL oxygen transfer coefficient (m/h)
K1 general biological decay rate
K2 temperature specific decay rate
L latitude of pond location (radians)
LIT net solar radiation penetrating water (ly/m)
LITDT extra-terrestrial solar radiation flux (ly/h)
OPH light intensity level producing photoinhibition
OS oxygen solubility at given temperature (mg/liter)
OP pond oxygen level (mg/liter)
P instantaneous photosynthetic rate
PHY phytoplankton biomass (rag/liter as carbon)
PMA X maximum photosynthesis rate
R radius vector (dimensionless)
RE reaeration rate (mg oxygen/h)
REF reflectivity of water surface (dimensionless)
SC solar constant (1.94 ly/min)
SR hour angle of sunrise (h)
TMP temperature modifier factor
W wind speed (m/s)
WTMP water temperature (°C)

INTRODUCTION

A basic task of pond management for fish culture is the maintenance of dissolved
oxygen at a level suitable for fish survival and growth. Pond managers can monitor
the dissolved oxygen level at a given point in time, but predicting potential critical
periods of low concentration is difficult. A few researchers have developed empirical
models to predict early morning pond dissolved oxygen concentration, given an initial
 COMPUTER MODEL OF OXYGEN BEHAVIOR IN A Q U A C U LTU R E POND 247

set of conditions (Busch et al., 1977; Boyd et al., 1978; Boyd, 1979). These models
have lumped many changes occurring in physical and biological processes by using
statistical techniques such as multiple regression, or by making simplifying assump-
tions. The models have been tested and used specifically for catfish ponds in Auburn,
Alabama. In order to gain a better understanding of events which can contribute
to low dissolved oxygen concentration, such as pond destratification and heavy
phytoplankton blooms, a model based upon known physical, chemical, and biological
mechanisms has been created and is presented here. This model simulates the hourly
fluctuation of dissolved oxygen in an aquaculture pond over a 24 h period, as
influenced by the consumption and production of oxygen by phytoplankton, fish,
detritus and reaeration. The model is neither site nor species specific and input para-
meters can be adjusted to accommodate most pond conditions. The model was
developed with three primary objectives in mind:
1. The establishment of a conceptual framework which unifies theories from
several disciplines.
2. The determination of components which have the greatest effect on dissolved
oxygen.
3. The identification of areas of research which could improve pond management.

THE MODEL

The relationships between the components, described quantitatively by the dissolved

oxygen flux model, are represented in Fig. 1. Wind, solar radiation and fish feed are
the major components which directly affect the rates of biological and physical
processes influencing pond water dissolved oxygen (DO) concentration. The solar
radiation at the surface of the water attenuates through the water column. The effec-
tive light intensity in the water column directly affects the phytoplankton population
which in turn, increases DO during the day via photosynthesis, and utilizes oxygen at
night through respiration. Decaying phytoplankton, unconsumed fish feed, and fish
waste products also decrease DO as represented by the sediment oxygen demand. As
the wind agitates the water surface, it influences the oxygen through reaeration. The
fish will also require oxygen for respiration, and will increase the DO demand. Equa-
tions describing each of these processes will be presented in the following sections.
Pond model equations are tied together by a mass balance, where the sum of the
reactants and inflow concentrations equals the sum of the outflow and accumulated
concentrations. A non-steady state approach is utilized to account for changing
environmental inputs. Values of rate coefficients used in the mass balance equations
are specified by the program or calculated hourly over a 24 h period. Each major
component influencing dissolved oxygen is presented in a modular structure and
evaluated independently of, or in relation to, other components.
248 D.I. MEYER,D. E. BRUNE

Reoerotion
FISH ~ SOLAR
RADIATION

Pholosynfhesis ~/?/

( ~.f,.~!,/~,,'~ - MesP' ron° n \ Respiration/

~ Waste \Sedlmenfaxygen / ,Algae .

Fee°,o..e. \pro,oc,ion \°ema°.
///~///~///~///~///--~///~///~//~, ///~///~--///~///~///
SEDIMENT

Fig. 1. Pictorial representation of pond oxygen flux model.

Solar radiation
The total solar energy reaching the earth's surface is the sum of direct solar radia-
tion and scattered or diffuse radiation. The amount of radiation per unit time depends
on the solar constant (1.94 ly/min), the latitude of the location (L, degrees), the time
of the year (JD, Julian date), the influence of the atmosphere (ATM), the albedo of
the earth's surface (REF) and the elevation of the location. A Tennessee Valley
Authority simulation study on solar radiation (1972), adapted by Orlob (1979), has
been implemented in the model.
The elevation of the location is assumed to be at sea level and the degree of obstruc-
tion between the location and the horizon are assumed to be negligible. The declina-
tion of the sun is given as if the observation was made at the center of the earth.
Atmospheric effects; reflectivity of the pond, latitude of the pond site and degree of
cloudiness (CL, 0 < CL < 1) affect the incident radiation, and can be indicated by the
user.
Daily, the radius vector (R, eqn (1)) and the declination of the sun (December,
eqn (2)) are calculated, given the Julian date and the latitude of the pond site. Times
of sunrise (SR) and sunset are calculated (eqn (3)) from the declination, radius vector,
and Julian date. The rate of solar radiation for each hour of daylight is calculated
(LITDT, ly/h, eqn (4)) with the maximum rate occurring close to hour 12. The
incident solar radiation at the water surface (LIT) is calculated using eqn (5). These
equations are as follows:
 C O M P U T E R M O D E L O F O X Y G E N B E H A V I O R IN A Q U A C U L T U R E POND 249

R = 1 + 0.17(cos (2zr/365)(186 --JD)) (1)

DEC = 23.45 0r/180)(cos (27r/365)(172 -- JD)) (2)
SR = (12/~) arccos ((tan L) (tanDEC)) (3)
L I T D T = 60SC[((DT + 1 -- SR)(siu L)(sin DEC)
+ (12u/12(cos L)(cos DEC))((sin u/12)(DT + 13)
x (sin 7r/12)(Dr+ 12))]/R 2 (4)
L I T = LITD T(A TM) ( 1 -- R E F ) ( 1 -- 0.65 CL 2)/60 (5)

Light attenuation
Light intensity within the water column can be described by the Beer-Lambert
Law, where the light intensity is attenuated exponentially with depth (D; Warren,
1971). The light extinction coefficient (EC) is influenced by the absorption and
scattering of light within the water column caused by water itself, and by the dissolved
and suspended substances of biological and non-biological origin (Jerlov, 1968).
In modeling light attenuation, spectral modification below the top meter of light
penetration is sufficiently small for the mean vertical extinction coefficient of white
light to be used (Hutchison, 1957). The extinction coefficient and distribution of each
substance in the pond is assumed completely mixed throughout the euphotic zone and
vertical migration of algae or consumers within the mixed layer are ignored. Only 50%
of the total light spectrum is available for photosynthesis (Strickland and Parsons,
1965). The effective light (ELIT) is calculated using eqn (6) below:
ELIT = 0.50(LIT) exp [--(EC)(D)] (6)

Reaeration
Depending on wind speed, pond oxygen concentration and water temperature,
oxygen will diffuse through the pond surface, tending to move in the direction which
will maintain the pond dissolved oxygen level at saturation. The overall oxygen
transfer coefficient is strongly dependent on the interface conditions. As wind speed
and surface water movement increase, potential convective oxygen transfer is
enhanced and the oxygen transfer coefficient increases many fold (Odum, 1956). An
empirical relationship between wind speed (W, m/s) and the oxygen transfer coeffi-
cient (KL, m/h) was established by Banks and Herrera (1977) (eqn (7)):
KL = 0-0036(843W °'s -- 3-67 W + 0.43W 2) (7)
Given that the portion of the water column under consideration is well mixed, the
contribution of reaeration to the pond DO level can be described as follows:
R E = IO00(KL )(AR )(OS -- OP) (8)
250 D. I. M E Y E R , D. E. B R U N E

Reaeration (RE, mg oxygen/h) is a function of the oxygen transfer coefficient, the

surface area of the pond (AR, m2), the pond dissolved oxygen level (OP, mg/liter) and
the level of dissolved oxygen the pond will contain at saturation (OS, mg/liter).

Phy toplankton
Factors influencing gross phytoplankton production rates include phytoplankton
density, species composition, age and physiological stage, as well as variations in
incident solar radiation, light attenuation in the water column, water temperature and
nutrient conditions. In modeling the effect of environmental factors on the rate of
photosynthesis, one of two mathematical functions may be chosen. For example,
depending on the effective light intensity, the photoplankton may be light limited or
photoinhibited; the proper response is selected and the average rate of photosynthesis
over a specified depth range is calculated. Given the average rate of photosynthesis, the
rate of oxygen production due to photosynthesis and the biomass of fixed phyto-
plankton is calculated. The contribution of phytoplankton to the pond DO level is
the difference between oxygen evolved through photosynthesis and that oxygen used
for respiration. First, the two major equations modeling photosynthesis are described,
including temperature and light effects. Phytoplankton respiration follows photo-
synthesis. The equations are used to simulate the rate of photosynthesis (P, h-~).
Smith's eqn (10) is used to simulate productivity at low light intensities where the
photosynthetic rate depends linearly upon the light intensity (Smith, 1936):

(ISL)(ELIT)
P= [ (ISL)(ELIT) ]o.s (10)
1 + [(PMAX)(TMP) 2J
Steele's eqn (11) is used at light intensities greater than the value where the photo-
synthetic rate becomes light-saturated (Steele, 1962):

(ELIT)(PMAX )( TMP)
e= (11)
exp 1
Steele's equation includes photoinhibition effects at high light intensities. Neces-
sary parameters for eqns (10) and (11) include the light extinction coefficient (EXT,
m-l), solar radiation intensity at a given time (LIT, ly/min), and the temperature
modifier (TMP, O<TMP< 1) and user-specified inputs which are species specific
(maximum photosynthetic rate (PMAX, h-l), initial slope of the photosynthesis versus
light intensity curve (ISL, 1/h/ly/min), light intensity occurring at the onset of photo-
inhibition (OPH, ly/min) and boundaries of the water column under consideration
(D 1, D2, m)).
 C O M P U T E R M O D E L O F O X Y G E N B E H A V I O R IN A Q U A C U L T U R E POND 251

This average photosynthetic rate (AP, h -1) for a simulated pond section is:

A P - (D2 Dr) PdD (1:)

1

(a) Temperature effects. The rate of photosynthesis exhibits a non-linear response

to water temperature, and the response curve is skewed to the right (Thornton and
Lessem, 1978). The maximum temperature and optimal temperature range have been
used as parameters to modify the curve to meet the temperature criteria for a given
dominant phytoplankton species. A temperature multiplier, a dimensionless number
between 0 and 1, modifies the rate of photosynthesis. Under optimal conditions,
temperature does not inhibit photosynthesis and the multiplier equals 1. The multi-
plier diminishes in value as the water temperature moves away from the optimal
temperature. The maximum rate of photosynthesis is multiplied by the temperature
multiplier to model this major effect of the actual rate of photosynthesis for a given
time period in eqns (10) and (11).

(b) Light efJbcts. Light is another important factor affecting photosynthesis. At

low light intensities the absorption of light energy limits the rate of photosynthesis,
independent of temperature. The rate of photosynthesis exhibits a linear response to
light intensity, proportional to the light absorbed by the photosynthetic pigments.
After reaching a species-specified light level, the rate of photochemical processes
exceeds the rate of enzymatic reactions and the rate of photosynthesis is no longer
light-limited. The maximum rate of photosynthesis, given non-light limiting or light
inhibiting conditions, is limited by the water temperature. In exceedingly bright light,
the rate of photosynthesis decreases as a result of the photooxidation of critical
enzymes and the inactivation of chlorophyll (Groden, 1977). Given the initial slope of
the linear response of the photosynthesis rate to light intensity, the maximum rate of
photosynthesis and the light intensity at the onset of photoinhibition, a curve may be
described reflecting the rate of photosynthesis against light intensity. This function is
assumed to be constant over depth and time, and is incorporated into eqn (10).

(c) Phytoplankton respiration. Using laboratory freshwater culture systems,

respiration rates have been recorded in response to photosynthesis, phytoplankton
species, water temperature and cellular stores. Endogenous respiration, the energy
required to maintain a certain level of biomass, increases exponentially with tempera-
ture (WTMP, °C) (according to Riley (1946) and Aruga (1965)). Jewell and McCarty
(1971) measured the initial respiration rate in their study dealing with the rate and
extent of algal decomposition. Using the exponential function by Riley (1956) and the
252 D.I. MEYER, D. E. BRUNE

initial respiration by Jewell and McCarty (1971), the endogeneous respiration rate
(ER) for the phytoplankton biomass (PHY, mg/liter) can be calculated (eqn (15)):
ER = 0.0012(PHY) exp (0-069) (wrMe) (15)
Photorespiration, occurring during photosynthesis, is used i~o oxidize organic
components, and is assumed to be proportional to gross photosynthesis. Endogenous
respiration and photorespiration are additive in this model.
In the present study, adjusted first-order decay rates were used to represent oxygen
uptake by detrital decay. The rate of decay and biological oxygen demand (BOD)
required for decay is determined by the type of matter decaying, the medium in which
the decay takes place and the temperature of the medium. In assessing the biological
oxygen demand at time T (BODT) of each component for an aquaculture system, the
following factors have been estimated: the quantity of decaying material present; its
ultimate BOD (BODL, mg O2/mg material) and the rate of decay (K, h-l); and the
water temperature (WTMP, °C). Detrital components include decaying phytoplankton,
unconsumed fish feed and fish waste products.
The decay rate constant is altered (K2, h-l), to account for pond water tempera-
ture, in eqn (13). The general relationship describing first-order decay over time (T,
days) is given in eqn (14):

K2 = KI(1 "056) WTMP-20 (13)

BODT = BODL(1 -- exp (-- T(K))) (14)

Fish respiration
The oxygen consumption of fish is affected by many factors, including water
temperature and oxygen content, carbon dioxide level, size of fish, activity and photo-
period. Using respiration flasks, researchers have been able to establish relationships
among various factors, and minimum dissolved oxygen requirements are known for
many fish species (Davis, 1975; Fry, 1971). Catfish were used as a test species for the
pond model. Andrews and Matsuda (1975) studied the effect of water temperature,
fish weight and dissolved oxygen on fish respiration; the catfish were stocked in 2-5-m
diameter culture tanks to simulate pond conditions. Using these data from this
experiment, Boyd (1979) performed a multiple regression and developed an equation
for catfish respiration (FR) using fish weight (FW, g), water temperature (WTMP) and
dissolved oxygen (OP, mg 02) as variables, combining the effects of fed and unfed fish.
These equations are:
FR = 10x (16)
where:
x = (--0-999) -- (0-000957)(FW) + (6 x 10-7)(FW) 2 + (O.0327)(WTMP)
-- (8.7 x I O-6)(WTMP)2 + (3 × I O-v)(WTMP)(FW)
 C O M P U T E R M O D E L O F O X Y G E N B E H A V I O R IN A Q U A C U L T U R E POND 253

and:
(FRO)(OP)
FR - if DO < 7.0
7

FR = (FRO) if DO > 7.0

RESULTS

When developing the DO flux model, both a component sensitivity analysis and a field
data verification study were completed. Validity of assumptions and constructs used in
model development lead to an analysis of areas of emphasis for the development of
a more comprehensive analysis of DO fluctuations, as described in the component
evaluation section.

Sensitivity analysis
Holding other parameters constant, the level of cloudiness was varied. As the level
of cloudiness increased, the pond DO decreased. A total cloud cover reduced the pond
dissolved oxygen by as much as 2 mg Oz/liter as compared to pond DO on a clear,
sunny day. The difference demonstrated the effect of solar radiation on net oxygen
production by phytoplankton (Fig. 2). The DO trend is similar to that produced in
a study on solar radiation effects (Boyd et al., 1978).
A second set of simulation runs were performed to determine how much influence
wind would have on the pond DO at different depths, given reaeration as the sole
contributor to pond DO (Fig. 3). As the wind speed increased, the time elapsed before
reaching saturation decreased. In all cases, the rate of aeration was greatest when pond
DO was furthest from saturation. The effect of the reaeration rate was found to be less
significant as pond depth was increased.
A third set of simulation runs looked at component sensitivity, the relative effects
of phytoplankton biomass, wind speed and BOD on pond oxygen conditions over a
period of 24 h. These three components appeared to have the greatest effect on pond
DO; specific tests and results are described in detail by Meyer (1980).

Field verification
In his study of metabolic rates of pond ecosystems under intensive catfish cultiva-
tion, Mezainis (1977) monitored the dissolved oxygen levels at 0-3 m intervals of
depth every 3 h, for a 24 h period. Concurrent chlorophyll measurements were by
Augusto (1975). The catfish pond was located at the Auburn University Experimental
Station, Alabama, and had an average depth of 1-5 m. Using the data collected by
Mezainis and Augusto, dissolved oxygen levels from the months of April and November
were simulated. During April, the pond was stratified, and the upper mixed layer (to
0.6 m) was simulated. During November the pond was totally mixed and the entire
254 13. I. M E Y E R , D. E. B R U N E

CL= 0

,_I
w
bJ
.J C L - - 1.0
Z
w
o
x
0
c~
w
>,
0
(t)

4 I I
800 1600
TIME OF DAY (HOURS)
l:ig. 2. The effect of cloudiness on pond oxygen levels.

water column was simulated (to 1.5 m). Data on secchi disk measurements and BOD
measurements were incomplete, and a few input parameters were estimated.
Figure 4 illustrates that the computer model is able to simulate the field data for
April. The solid line represents the results from the computer simulation. The dashed
lines bound the range of values at sampling stations in the pond. In the morning hours
the pond dissolved oxygen is failing due to the p h y t o p l a n k t o n and fish respiration and
detrital decay. Following sunrise, the pond DO concentration increases as the phyto-
plankton produce more oxygen through photosynthesis than is consumed through
respiration and decay. The phytoplankton photosynthesis decreases as the intensity o f
the solar radiation decreases in the late afternoon. November results (Fig. 5) follow
the basic trend o f a 24 h oxygen flux as reported by Mezainis, but are not as accurate
 COMPUTER MODEL OF OXYGEN BEHAVIOR IN AQUACULTURE POND 255

1
l m depth
~ 2 m depth

11 rnls
/ I
lob
E
/
-8 / / 6 m/s
-J
IJJ / /
IJJ
_J /
/
Z
taJ /
>-
x
0 J 1 mls
r,,
,,,7
/ j
/
0

/
/ j J
/

I I I I I

10 20 30 40 50
ELAPSED TIME (HOURS)
Fig. 3. The effect of wind speed on pond oxygen levels.

at representing the data. This discrepancy is likely due to the fact that the pond may
not have been as well mixed during this time period as previously suggested by
Mezainis.

Component evaluation
(a) Solar radiation. The set of equations describing solar radiation (eqns (1)-(5))
are based on a sound theoretical and experimental basis, and have been applied to
solar radiation modeling studies. The solar radiation model is adequate, especially for
its intended use in this model. Its importance lies in the fact that solar radiation is
a forcing function which allows flexibility in analyzing pond conditions throughout
the world.
256 D. I. M E Y E R , D. E. B R U N E

A
• \
-- I \

E I ",
I

--I
ILl -- III
>
,,,.
6
/ / \
Z ¢ r --/.._ \
tu ," / / -~" . . . . \
~" jl /

a
..... ,,,, ,,/ \
I.ul ~ "-~,' ,1/
0
""--..X J'f
"/
_m
o 4

..... 9 5 % Confidence infervols, pond s-14

- - Compufer simulofion

0 400 1200 2000

TIME OF DAY (HOURS)
Fig. 4. A comparison of field data collected by Mezainis (for the month of April) to computer
simulation.

(b) Attenuation o f solar radiation. The major factor limiting the accuracy o f the
light attenuation function is the extinction coefficient. Using a relationship developed
by Poole and Atkins (1929), the secchi disk reading is used to determine the extinction
coefficient. The components affecting the secchi disk reading are not comparable from
one pond to another (Steeman Nielsen, 1975). For example, turbidity caused by the
presence of clay particles may alter the light extinction coefficient of one pond;
a phytoplankton population may alter the light extinction coefficient in another pond.
Short-term answers in any one pond may be handled using secchi readings because
these components may not vary widely with a single location. For research and a
general comparison, further experimentation on factors affecting the extinction
coefficient is necessary. Early field studies by Riley (1956), Lorenzen (1972) and
 COMPUTER MODEL OF OXYGEN BEHAVIOR IN AQUACULTURE POND 257

10-

t~n m S, %%%%%
E ~,S
-.I
1.1.1
I.fl
._1 "... , SS //
z
LLI
%. ....... " / I
>-
X
0
", /
a
LIJ
~ sJ
0
7 - "'.,/

9 5 0 Confidence intervols, pond s-14

Computer simulotion

6__/1, I I I
0 400 1200 2000
TIME OF DAY (HOURS)
Fig. 5. A comparison of field data collected by Mezainis (for the month of November) to
computer simulation.

Sakamoto (1966) established empirical relationships between the extinction coeffi-

cient and factors such as chlorophyll A and inorganic and organic matter. Many studies
have limited application because the light comparison depth and chlorophyll A con-
centration have to be measured before the influence of the extinction factors can be
calculated.

(c) Reaeration. The relationship used to describe the reaeration coefficient is based
on data from large, well-mixed bodies of water. Direct measurements of reaeration in
aquaculture ponds (smaller water bodies) is not available. The influence o f the wind
stirring up sediments at the bottom o f ponds also needs further investigation.
258 D . I . M E Y E R , D. E. B R U N E

(d) Phytoplankton. The equations describing phytoplankton photosynthesis and

growth responses to temperature and light use the current state of the art approach.
Unfortunately studies addressing specific occurrences on an hourly basis over a 24 h
period are scarce and experimental data is difficult to collect and interpret. Factors in
need of further development include phytoplankton production and sink rates. For
example, phytoplankton respiration rates may increase exponentially at higher
temperatures. In the lower range of temperatures a linear approximation may be more
realistic (Canale et al., 1976). Presently, an exponential response curve has been
incorporated into the DO flux model.
In a simulation study by Groden (1977) both the maximum temperature, initial
slope and onset of inhibition of the phytoplankton-light curve are calculated, given the
water temperature, optimum temperature and maximum rate of photosynthesis.
Applicability of the relationship is unknown and access to data that must be supplied
by the user is limited. However, composite values available for various phytoplankton
species throughout the world and in laboratory studies are summarized by Parsons
et al. (1977).
Algae respond to the daily cycle of light intensity and will reach their maximum
rate of photosynthesis at a light intensity which is a function of the daily solar radia-
tion (Ichimura, 1960). Seasonal solar radiation also affects phytoplankton populations.
As seasonal solar radiation becomes pronounced, cellular concentration of chlorophyll
will vary affecting the chlorophyll to carbon ratio; a ratio which must be used to
convert field data to a model input parameter. This ratio can be set by the user to be
season-specific representing the time of year under study.
The phytoplankton production rate is based solely on the rate of photosynthesis
and incorporates two user-specified ratios: chlorophyll A-to-algae and oxygen-to-
algae. Varying either will effect the DO flux. The first one is used when interpreting
field data on phytoplankton biomass, generally reported as mg chlorophyll A/rag °C h.
Chlorophyll A content varies with the light intensity to which the phytoplankton are
adapted; it also varies among species. The oxygen-to-algae ratio is based on a thermo-
dynamic analysis of photosynthesis as presented by Hendricks and Pote (1974).
The algal sink rate is also a user-specified coefficient. Algal decay is influenced by
cell size, environmental conditions, and the physiological condition of the algae. A few
of these factors have been described quantitatively in models by DePinto et al. (1976)
and Canale et al. (1976). The importance and accuracy of the mathematical repre-
sentation has not been evaluated in pond systems.

(e) Sediment oxygen demand. The production of decaying material, the ultimate
BOD of that material and its rate of decay determine the BOD of the detritus. Accord-
ing to Scavia (1979), first-order rates represent actual data better than constant coeffi-
cients although the decay rate function has not been quantitatively validated through
field and laboratory studies. Sources of BOD included in the DO flux model include
fish manure, unconsumed fish feed and phytoplankton decay. The ultimate BOD and
 C O M P U T E R M O D E L O F O X Y G E N B E H A V I O R IN A Q U A C U L T U R E POND 259

decay rate of each of these sources are not well-defined and need to be clarified
through experiments. Another option would be to approach the question of detrital
BOD by simulating the bacterial population fluctuations instead of assuming that the
rate of decay responded only to temperature. The presence of bacteria active in
detrital decay processes is implicit in the decay function.
The model does not account for the location of the detrital material in the water
column, as would be required to stimulate a stratified pond or an older pond with
heavy sediment build-up.

DISCUSSION

The model as presented utilizes basic principles concerning oxygen transfer, biological
oxygen production and consumption and degradation of organic matter. It can be used
with experimental data and in hypothesis testing. Its construction and operation has
elucidated key areas for future research, with the goal of increasing its applicability for
aquaculturists, and developing a cohesive model of short term oxygen production as
a tool for scientific study. The major constructs in the model have been formalized in
other studies of aquatic systems in limnology and aerobic waste treatment ponds in
sanitary engineering. This model is an incorporation and synthesis of these constructs,
applicable to aquaculture ponds. Areas where available information is tentative have
become apparent, as evaluated in the previous section. A short term model emphasizes
changes occurring in short time periods minimizing the seasonal effects of changing
composition of phytoplankton species, fish oxygen requirements, sediment build-up
and water balance.
A major goal is to simulate pond DO during infrequent, yet critical events, such as
cloudy days in the summer, blue-green algae die-offs and windy or still days. One
condition which cannot be simulated presently is pond destratification from wind and
temperature changes. The assumption that the horizontal sector being simulated is well
mixed limits the representation of this condition, because influence from other hori-
zontal sectors is ignored. Modeling a stratified pond as a set of horizontal sectors,
where each sector is assumed well mixed and where one sector may influence the other,
may contribute to an accurate representation of this system. Mixing of water in a pond
is caused by various advection, dispersion and diffusion processes as well as circulation
patterns. The turbulence may stir up and aerate the suspended materials near the sedi-
ment-water interface increasing pond BOD and enhancing light attenuation. With the
incorporation of a hydrodynamic or mixing submodel, the following questions may be
addressed: Is the sediment layer often resuspended in a pond, aerating the benthos?
How does the circulation of a pond vary with its depth? How do various storm events
of known hydrodynamic characteristics affect a pond? Models of wind-generated
circulation have been reviewed by Banks (1975).
260 D.I. MEYER, D.E. BRUNE

When evaluated as a t o o l to be used by the fish farmer, the D O flux m o d e l is in

a d e v e l o p m e n t a l stage. The large n u m b e r o f input parameters c o n t r i b u t e to the flexi-
bility o f p o n d conditions, but m a k e it c u m b e r s o m e to use as a daily indicator. How-
ever, most inputs will not vary within a p o n d and could be set as constant input
values to d e t e r m i n e c o n d i t i o n s which could lead to critical D O levels. Implicit in the
use o f this m o d e l is a need for m o r e extensive field verification. As a research tool,
the m o d e l has imposed a m a t h e m a t i c a l f r a m e w o r k for current theories applicable to
D O flux. Given that B O D , p h y t o p l a n k t o n and wind speed have a major effect on the
p o n d DO. L a b o r a t o r y studies which will support a better understanding o f these
c o m p o n e n t s is r e c o m m e n d e d .

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