Aquacultural Engineering 34 (2006) 214–223

www.elsevier.com/locate/aqua-online

Linear versus Monod representation of ammonia oxidation

rates in oligotrophic recirculating aquaculture systems
Ronald F. Malone a,*, Jon Bergeron b, Chad M. Cristina c
a
Department of Civil and Environmental Engineering, Louisiana State University,
Baton Rouge, LA 70803, USA
b
U.S. Public Health Service, 4060 Wheaton Way Ste. E., Bremerton, WA 98310, USA
c
Department of Civil Engineering, University of South Alabama, Mobile, AL 36688, USA

Abstract

Monod kinetics are widely used to model nitrifying biofilters. However, these kinetics are incapable of representing the
collapse of volumetric TAN conversion rate (VTR) under high organic loadings. Failure to recognize the underlying
heterotrophic interference can lead to calibration issues as a single Monod function is applied across contrasting levels of
carbon loading. This, plus an historic bias towards the analysis of peak carrying capacities leave modelers poorly prepared to
serve the needs of a mariculture industry demanding oligotrophic designs for broodstock maturation and larval/fingerling
production. Consequently, data was generated by a Monte Carlo technique under the assumption of heterotrophic inhibition to
nitrification. The data was used to compare the accuracy of calibration of the Monod relationship using the traditional
Lineweaver–Burke and Eadie–Hofstee calibration methods against direct linear regression for low substrate (mesotrophic/
oligotrophic) regimes. The results indicate that a simple linear relationship with a zero intercept, calibrated on data ranging from
0.1 to 0.5 g-TAN m
3, is most suitable for the representation of the mesotrophic/oligotrophic performance of nitrifying biofilters
based on a comparison of SSE for both the Monte Carlo and field data analyzed herein. Additionally, the coefficient of variation
was found to be between 7 and 8% for the parameter t, which is the slope of the linear relationship between total ammonia
nitrogen (TAN) and VTR while the CV for the Monod parameters ranged between 22 and 143% for VTRmax and between 29 and
137% for the apparent half-saturation constant showing the improved stability of the linear model to that of the Monod model.
# 2005 Published by Elsevier B.V.

Keywords: Monod model; Broodstock; Oligotrophic; Modeling; Inhibition; Nitrification kinetics; Linear regression

1. Introduction Consequently, these systems are increasingly being

The use of recirculating systems for the condition-
ing of marine broodstock is gaining wide acceptance.
* Corresponding author.
E-mail address: RMalone@lsu.edu (R.F. Malone).
used to produce larvae, fry and fingerlings for grow out
systems (Malone, 2002). Although this increase in
utilization appears to have stemmed from the need for
precise control of temperature and light regimes,
adoption now is often driven by the desire to precisely
control water quality and nutritional conditions in the

0144-8609/$ – see front matter # 2005 Published by Elsevier B.V.

doi:10.1016/j.aquaeng.2005.08.005
 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223
sensitive, early stages of life. Also the isolated nature
of a closed recirculating system provides benefits in
the control of diseases that have historically deva-
stated the marine industry.
Most marine hatchery systems require oligotrophic
water quality conditions in order to maintain a healthy
stock. Broodstock systems generally require the total
ammonia nitrogen (TAN) concentration to be in the
range of 0.1–0.3 g-N m
3. Larval systems typically
require TAN concentrations less than 0.1 g-N m
3 and
fingerlings typically require a TAN concentration less
than 0.5 g-N m
3 (Malone and Beecher, 2000).
Substrate utilization can be divided into the
following three processes (Zhang and Bishop,
1994). First, the substrate must diffuse from the bulk
fluid, through a static boundary layer, to the biofilm
surface. Under oligotrophic water quality conditions,
low substrate concentrations reduce the concentration
gradient between the bulk fluid and the static boundary
layer, thus limiting the areal nitrification capacities of
biofilters and dramatically increasing their expense. In
the second process, substrate must diffuse through the
biofilm to the embedded bacteria. In the oligotrophic
water quality conditions, ammonia diffuses into
a relatively thin, vertically homogenous biofilm
that is dominated by autotrophs due to the low
biochemical oxygen demand (BOD) of the culture
water (BOD < 5 g m
3 Malone and DeLosReyes,
1997). Eutrophic conditions create diffusion limita-
tions within the biofilm itself. Biofilms under
these water quality conditions are characterized by
layered biofilms that are dominated by relatively fast-
growing heterotrophs due to the higher BOD
(BOD > 20 g m
3, Malone and DeLosReyes, 1997).
Ammonia must diffuse through the heterotrophic layer
of the biofilm to reach the nitrifiers below the surface.
Finally, once the substrate reaches the bacteria within
the biofilm, these bacteria can consume the substrate.
While sizing tables for oligotrophic conditions are
known to be approximate (Malone and Beecher,
2000), the use of computer models to simulate the
nitrification behavior of a variety of biofilters has been
met with some success (Golz et al., 1999). A multi-
compartment computer model was recently developed
to facilitate the design of platform-based broodstock
and fingerling production systems to be located off the
coast of California. Compound Monod models
reflected the impact of ammonia, oxygen and
215
alkalinity on the TAN and nitrite-N conversion
capacities of the biofilters. Although approximate in
nature, the model proved helpful in the sizing of bead
filter and fluidized bed combinations as well as in
determining pumping and heating capacities needed to
support the platform activities.
Certainly, our understanding of nitrification
kinetics has advanced to the point that these models
are useful in a pragmatic sense both as analysis and
communication tools. However, these same applica-
tions tend to quickly reveal the shortcomings of our
current knowledge and engineering practice, since
small variations in model parameters may have
significant effects on the projected filter size for a
given application. The proximate nature of the Monod
relationships that are used to represent the fixed-film
nitrification kinetics reflects the complexities of
transport and enzymatic reactions that take place in
biofilters (Bishop, 1997). The core theory is rooted in
Michaelis–Menton enzyme kinetics, which can be
simply represented as:
Vmax S
Rate ¼ (1)
kM þ S
where Vmax is the maximum specific rate of substrate
utilization (g day
1), S the substrate concentration
(g m
3) and kM is the half-saturation constant
(g m
3). Eq. (1) is applicable only to conditions of
constant enzyme availability. This relationship is often
applied to bioreactors exhibiting dynamic bacterial
growth in the Monod formulation:
Kmax XS
Rate ¼ Vb (2)
kM þ S
where Kmax is the biomass normalized maximum rate
constant in (g of S converted) (g biomass)
1 day
1, X
is the bacterial biomass (g biomass m
3) and Vb is the
reactor volume (m3).
The volumetric TAN conversion rate (VTR) has
been similarly modeled as (Malone et al., 1993):
Rate VTRmax A
VTR ¼ ¼ (3)
Vb kb þ A
where VTR is the volumetric TAN conversion rate
(g m
3 day
1), VTRmax the maximum VTR attainable
(g m
3 day
1), A the TAN concentration in the bulk
fluid (g m
3) and kb is the apparent half-saturation
constant (g m
3). Eq. (3) inherently presumes that the
216 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223
bacterial mass (and in turn the enzyme mass) is
proportional to the substrate under conditions of
steady loading (on a time frame of days). Additionally,
in fixed-film biofilters, it has been found that the half-
saturation constant is not a universal constant. Rather,
the value is constant only for a specific set of hydraulic
conductivity, biofilm thickness and interstitial velocity
conditions and can change with filter management
procedures (Malone et al., 1993).
It is perhaps just fortunate that surface area
dependent diffusion reactions are also well repre-
sented by Monod formats. The importance of diffusion
into biofilms has not only been elegantly recognized
by early kinetic modelers (Harremoes, 1982), but now
has been verified by direct measurements within
biofilms (Zhang and Bishop, 1994; Zhang et al., 1995;
de Beer et al., 1997). Data is readily available to
calibrate Eq. (3), since VTR is easily calculated by
flow and concentration data (Malone and Beecher,
2000).
Eq. (3) is transformed using either the Eadie–
Hofstee or the Lineweaver–Burke method. Parameter
values for VTRmax and kb are determined through a
linear regression analysis. The linearized versions of
Eq. (3) using the Eadie–Hofstee and Lineweaver–
Burke methods can be found in Eqs. (4) and (5),
respectively, as:
VTR
VTR ¼
kb þ VTRmax (4)
A waters. Satoh et al. (2000) used microelectrodes to
1 kb 1 1
¼ þ (5)
VTR VTRmax A VTRmax
Both methods of linearization have their limita-
tions. The Eadie–Hofstee method incorporates VTR
into both the ordinate and abscissa in the regression
and is thus more susceptible to measurement error
than is the Lineweaver–Burke method, which does not
include VTR in the ordinate. The Lineweaver–Burke
method uses reciprocals of measured values. If the
number of data points is spread evenly across the range
of interest, this method gives inordinate weight to
VTR values when A is less than 1 g m
3.
The historic emphasis of the recirculating com-
munity has been focused on freshwater grow out
systems with high substrate regimes as opposed to the
low substrate regimes required for hatcheries. This
emphasis resulted in databases with numerous TAN
measurements under high substrate loading conditions
and relatively few TAN measurements under low
substrate loading conditions. This bias has been
further exacerbated by the difficulties associated with
obtaining accurate TAN values below 0.3 g m
3. The
combination of these two biases tends to counteract
the limitation of the Lineweaver–Burke method and
yield coefficients that are biased in favor of the mid-
range of TAN values. The end result are calibrations
that are reasonably accurate in predicting filter
performance under regimes of high nitrogen and
organic loading, but, weak or simply erroneous, in the
oligotrophic zone. Despite the limitations of para-
meter determination, many researchers have shown
that, in a pragmatic sense, the functional relationship
found in Eq. (3) can reasonably represent the VTR of
biofilters over a wide range of TAN concentrations
(Malone et al., 1993; DeLosReyes and Lawson, 1996;
Zhu and Chen, 2001; Golz et al., 1999).
Eq. (3) does not reflect the impact of heterotrophic
bacteria, which is widely recognized to impair both
TAN and nitrite conversion as organic levels in the
system increase (Hockenbury and Leslie Grady, 1977;
Harremoes, 1982; Bovendeur et al., 1990; Zhang and
Bishop, 1994). The algorithms of this model projected
maximum TAN conversion capacities well beyond the
point of known filter failure. Several researchers have
found that VTRmax is not a constant, but is dependent
on the carbon to nitrogen (C/N) ratio within the culture
determine the effect of C/N on the spatial distributions
of ammonia-oxidizing bacteria. They found that as C/
N increased, NH4+ oxidation rates decreased and
proposed that interspecies competition between
heterotrophs and nitrifiers may have been the cause.
Both Ohashi et al. (1996) and FDZ-Polanco et al.
(2000) reached similar conclusions when studying
biological aerated filters. Both teams reported a
decreasing proportion of nitrifiers in the biofilms as
a function of increasing C/N. Zhu and Chen (2001)
reported a 70% decline in VTR when the C/N ratio
was increased to 1.0 as compared to a pure ammonia
feed (C/N = 0). Gupta and Gupta (2001) found that the
nitrification rate increased from 0.81 g-N m
2 day
1
with increasing nitrogen loading despite high organic
loading. However, they found that the rate of
nitrification decreased at a high load of 3.35 g-
N m
2 day
1 and 32 g-COD m
2 day
1 and proposed
 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223
that the reduction was the result of some form of
inhibition process. These findings indicate the need for
the use of some form of inhibition kinetics as opposed
to fundamental Michaelis–Menten kinetics when
modeling these systems.
The second shortcoming identified with the current
practice is perhaps an historical artifact of the
development of biofiltration in aquaculture, which
has focused more on the heavy production end of the
loading regime than on the oligotrophic. Traditionally,
our principle basis for comparison of filtration capacity
has been maximum ammonia conversion capacity. The
maximum conversion capacity was viewed as the
principle characteristic determining the filters value in
the search for a cost effective, recirculating grow out
strategy for commodity foodfish. The databases, and in
turn, our kinetic constants reflect this biased view
towards heavily loaded regimes. The increased
emphasis on the production of marine species has
shifted the demand on our biofilters as they are
increasingly used for breeding, larval and fingerling
production. Increasingly the relationship between
extremely low substrate levels and conversion capacity
determines the filters sizing, not its peak carrying
capacity. Appropriate determination of the half-satura-
tion constant rather than the maximum conversion
capacity becomes critical. The methodologies used for
estimating low substrate (<0.5 g-N m
3) conversion
capacities must be revised and statistical techniques that
do not slant curve-fitting exercises towards high
substrate concentrations should be identified.
It is proposed herein that linear regression can be
used, without data transformation, to model oligo-
trophic systems (Chitta, 1993). Although this method
is not ideal from a purely scientific standpoint, it
would have several strengths. First, it would serve as
an intermediate step to improve modeling accuracy in
oligotrophic systems until a consensus can be reached
on the proper form of inhibition-based nitrification
kinetics. Second, since this linear relationship would
only be applied for TAN concentrations in the range of
0.1–0.5 g m
3, data collection could be simplified
through the use of chemical feeds, since heterotrophic
interference would be minimal under such low
loadings. Third, it would improve the engineer’s
ability to rapidly size biofilters for oligotrophic
applications. Finally, it could provide a simple means
for potential users of a given biofilter to compare
217
differing technologies based on a single parameter, the
slope of the regression equation.
2. Data generation and analysis
The limitation of Eq. (3) is that a single equation is
calibrated across two dramatically different operating
regimes. In a mathematical sense, it is argued that
biofilter performance is more appropriately repre-
sented by a type of inhibition kinetics. To this end,
Eq. (6) is presented. However, it should be noted that
the purpose of Eq. (6) is explanatory in nature and is
not being promulgated as the final form of inhibition
kinetics in aquacultural engineering. It is solely being
used to highlight the error that can be introduced into
VTR modeling by fitting a Monod relationship to a
process whose underlying behavior is governed by
inhibition kinetics.
Vm A
VTR ¼ (6)
kI þ A þ cA2
where Vm is the analogous to VTRmax (g m
3 day
1),
kI the analogous to the apparent half-saturation con-
stant kb (g m
3) and c is the constant (m3 g
1) used to
provide dimensional consistency.
Fig. 1 contains sample VTR plots comparing
Eqs. (3) and (6). The principle difference in the two
relationships is Eq. (6) ability to reflect the collapse of
nitrification capacity that is associated with increased
organic loading as opposed to the Michaelis–Menten
model whose VTR increases and approaches a
maximum as A approaches infinity. In this mathema-
tical configuration, A is used as a surrogate for organic
loading in the inhibition term, noting that the C/N
loading ratio remains fixed by the feed protein content.
If the focus is shifted from the entire graph to the
oligotrophic region (0 g m
3  TAN  0.5 g m
3), as
shown in the cut out of Fig. 1, we can notice the
discrepancy between the estimations of inhibition
kinetics through the choice of either Michaelis–
Menten kinetics or a simple linear regression between
A and VTR. Estimates of the required biofilter size
using these two different kinetic relationships can
differ by as much as 25%.
The first step in this analysis was data generation.
For this purpose, it was assumed that the inhibition
kinetics defined by Eq. (6) truly defined the
218 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223

Fig. 1. Sample plots comparing the estimation error in VTR when the estimation method is either a Monod kinetic model or a direct linear
regression. The inhibition curve represents ammonia oxidation within a biofilter for the given inhibition expression. The Monod and linear curves
represent best-fit estimates of the inhibition curve. Parameter values include Vm = 6000 g m
3 day
1, kI = 5 g m
3, c = 1 m3 g
1,
VTRmax = 1392 g m
3 day
1, kA = 1.231 g m
3 and t = 1075 day
1. It should be noted that the inhibition expression shown above is for
explanatory purposes.

conditional mean VTR. Eq. (6) produces a continuous
curve. However, it was necessary to introduce
variability into Eq. (6) to simulate field measurements
as:
 
Vm A Vm A
VTR ¼ N ; f
kI þ A þ cA2 kI þ A þ cA2
where N(Xbar, s) indicates a normal probability density
function with conditional mean Xbar and standard
deviation s. The constant, c, was defined as
1 m3 g
1. The constant f ( f > 0) represents the stan-
dard deviation as a fixed proportion of the conditional
mean. For the purposes of this exploration, f was set to
a value of 0.3. This value was chosen due to its ability
to create reasonable variability in the data while
minimizing the occurrence of negative values for
VTR, which was problematic with higher values of f.
The data were generated in the following manner.
N = 20 data sets were generated for each Vm–kI
(7)
combination including values of Vm equal to 6000 and
9000 g m
3 day
1 and kI = 5 and 10 g m
3. Four
values of VTR were randomly generated from Eq. (7)
for each value of A including A = 0.1 to 5.0 g-
TAN m
3 in increments of 0.1 g-TAN m
3. Four
values of VTR were generated to more closely
simulate field data where repeated values of A are not
uncommon. Furthermore, four values were found to
 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223 219

produce a manageable number of data points for
analysis. Fig. 2 contains sample plots of the Monte
Carlo data generated for each step of the analysis.
Once the data were generated from Eq. (7), two
methods of estimating the underlying function from
the generated data were attempted. The Lineweaver–
Burke (LB) method in Eq. (5) was used to fit Eq. (3) to
the generated data. The equation used to fit the direct
linear regression model was expressed as:
VTR ¼ mA þ b
The parameter m is the slope of the line relating
VTR and A while the parameter b is the y-intercept of
the equation.
Least squares analyses were performed to deter-
mine the values of the parameters m and b. When the
20 data sets were analyzed, it was found that b did not
statistically differ from 0 for any data set (P > 0.05).
This finding was not surprising since the function used
to generate the data had the property that VTR = 0
when A = 0.
This approach appears to contradict the concept of
Amin, the minimum, non-zero, concentration of A
required to sustain a steady state biofilm (Rittmann
and McCarty, 1980). However, the steady state
requirement for Amin is not met in this application.
Fixed-film biofilters are steady state on the order of
days but unsteady state on the order of hours due to
feeding. Consequently, values lower than Amin can be
sustained for limited durations between feedings, a
finding consistent with those of Furamai and Rittman
(1994).
The values of b in Eq. (8) were forced to zero and
(8)
new slopes, t, were computed. The new slopes were
used for the remainder of the analysis with the linear
relationship between VTR and A expressed as:
VTR ¼ tA (9)
where t is the normalized conversion capacity relating
VTR and A (day
1). Once the value of t has been
determined, the mass of TAN that a biofilter could
convert in one day could be determined.
Rate ¼ tAVb (10)
Under the typical conditions of application where
both the TAN loading (L) (g day
1) and the critical

Fig. 2. Sample plots for the Monte Carlo data generated in this analysis. Each set of data points represents only one out of twenty data sets
generated for each set of Vm and kI. In plot (A) Vm = 6000 g m
3 day
1, kI = 5 g m
3. In plot (B) Vm = 9000 g m
3 day
1, kI = 5 g m
3. In plot
(C) Vm = 6000 g m
3 day
1, kI = 10 g m
3. Finally, in plot (D) Vm = 9000 g m
3 day
1, kI = 10 g m
3.
220 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223
Table 1
Means and coefficient of variations (ratio of standard deviation to mean) for all estimated parameter values
Vm kI Mean estimated
(g m
3 day
1) (g m
3) VTRmax (g m
3 day
1) (CV%)
6000 5 1392 (22)
6000 10 1269 (20)
9000 5 2745 (143)
9000 10 3336 (137)
N = 20 Monte Carlo data sets were used for each Vm–kI pair to determine mean parameter values.
TAN concentration (AC) (g m
3) are user defined, the
filter size can be directly computed.
Lð1
Is Þ
Vb ¼
tAC
where Is is the in situ nitrification fraction (unit less)
representing that portion of the nitrification that occurs
on the tank and pipe walls (Malone and Beecher,
2000). Observed values of Is rarely fall below 30%,
however, this factor is sometimes neglected forcing a
an oversizing of the filter that contributes to the safety
factor of the design.
Parameter values for both the Monod and direct
linear regression (DLR) methods were estimated for
each of the N = 20 data sets and for each combination
of Vm and kI yielding a total of 80 parameter estimates.
There was one significant difference in this operation.
The Monod parameters were determined by lineariz-
ing the data across the range of 0.1–5.0 g m
3
(representative of typical practice) while the DLR
used only data in the 0.1–0.5 g m
3 range (as
proposed).
The mean parameters for each method and each
Vm–kI pair can be found in Table 1. The coefficient of
variation shows that parameter estimates for VTRmax
and kA were more variable than those of t. The
standard deviation for t was found to range between 7
and 8% of the mean value of t. However, the standard
deviation for VTRmax ranged from 22 to 143% of the
mean VTRmax and the standard deviation of kA ranged
from 29 to 137% of the mean kA. To further exacerbate
the problem, the Monod kinetic expression of Eq. (3)
contains two parameters, both of which are more
variable than the single-parameter linear model from
Eq. (9). The higher variability in the Monod kinetic
model creates uncertainty as to system performance
when these systems are modeled in the computer-
Mean estimated Mean estimated
kA (g m
3) (CV%) t (day
1) (CV%)
1 (48) 1075 (7)
2 (29) 561 (8)
1 (137) 1666 (8)
4 (121) 870 (7)
modeling environment. However, the relatively low
uncertainty for the DLR leads to greater stability in the
same situation.
Once the parameter values were determined for a
(11)
given data set, these parameters were used to
‘‘predict’’ the VTR within the 0.1–0.5 g m
3 range.
The sum of squared errors (SSE) was computed for
each data set and each kinetic model using only data in
this same range. Paired t-tests were then used to
determine if either method had a significantly lower
SSE than the other for a given Vm–kI combination. It
was found that the DLR had a significantly lower SSE
than the Monod model for each of the four data sets
(P < 0.05) verifying what was found in Table 1.
3. Comparison of methods using field data
Field data was sought to verify the results of the
Monte Carlo analysis. The data sets included two bead
filter data sets. The first set of data was collected by
Chitta (1993), while the second data set was collected
by Pfeiffer and Malone (2005). The data used in this
analysis can be found in Fig. 3 with the results of both
the DLR and Monod fitting methods found in Fig. 4.
Since data was not available values of TAN in excess
of 1.5 g-TAN m
3 for the Chitta (1993) data or above
0.8 g-TAN m
3 for the Pfeiffer and Malone (2005)
data, the Monod model was calibrated across the full
range of available TAN data for each of the two
available data sets. The DLR was calibrated only for
TAN values ranging between 0.0 and 0.5 g-TAN m
3,
consistent with the Monte Carlo analysis. The ability
for the model to mimic the data was tested, using SSE,
only for TAN values ranging between 0.0 and 0.5 g-
TAN m
3, which were again consistent with the
Monte Carlo analysis.
 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223 221

Fig. 3. Field data used to verify Monte Carlo analysis.

The SSE for Chitta (1993) was calculated to be
474,303 g2-N m
6 day
2 for the DLR and 508,660 g2-
N m
6 day
2 for the Monod kinetics for values of
TAN ranging from 0.0 to 0.5 g-TAN m
3 The data
from Pfeiffer and Malone (2005) resulted in SSEs for
the DLR and Monod of 3981 and 4280 g2-N m
6
day
2, respectively, for the same range of analysis. For
both data sets, the SSE of the DLR was found to be
lower than that of Monod kinetics. Clearly, this is not
an exhaustive analysis, though it does show the
potential of using a DLR to simplify modeling efforts
for oligotrophic water quality conditions.
4. Recommendations/suggestions
For the purposes of modeling VTR under
mesotrophic/oligotrophic water quality conditions,
where the TAN concentration must be kept below
0.5 g m
3, the direct linear regression model cali-
brated to localized data (TAN < 0.5 g m
3) was found
to provide significantly more accurate estimates of
VTR than Monod kinetics calibrated to the full data
range (TAN < 5.0 g m
3) based on an analysis of the
sum of squared errors in a Monte Carlo analysis
(P < 0.05). A similar analysis on field data also
indicated that the DLR yielded lower values of SSE
when estimating VTR for TAN values ranging from
0.0 to 0.5 g-TAN m
3. This observation should be
verified with analysis of additional field data from
oligotrophic/mesotrophic systems.
The estimated parameter values of the DLR were
found to be less variable than those of the Monod
kinetics based on the coefficients of variation. These
findings indicate that the use of the DLR may
introduce less error into the modeling process than
does Monod kinetics. Consequently, the proposed
premise seems to be valid. The use of the DLR instead
of Monod kinetics, under oligotrophic conditions,
could serve as an intermediate step to improve the

Fig. 4. Monod and direct linear regression fits of field data in the oligotrophic water quality range.
222 R.F. Malone et al. / Aquacultural Engineering 34 (2006) 214–223
accuracy of computer models used under these
conditions until a consensus can be reached on the
proper form of inhibition kinetics.
The use of t as a rating system would also be
valuable to engineers and consumers who are
designing or operating oligotrophic systems. These
filters could be compared by their t values from
Table 1. If we only look at the first two Vm–kI entries in
Table 1, we can refer to the first entry as having a t-
rating of 1075 day
1, based on the mean t found in
column 5, while the second entry would be referred to
as having a t-rating of only 561 day
1. Immediately, it
would be known that the filter rated at 1075 day
1
filter was more efficient on a media-volume basis than
the filter rated at 561 day
1 and would require a filter
comprising only half the volume of the lower rated
filter.
It is clear that new forms of nitrification kinetics need
to be investigated in order to represent filter collapse
under high organic loadings. These new kinetics will
likely resemble the inhibition kinetics presented herein
by including some non-linear term within the expres-
sion. However, it is unclear if fitting an equation across a
broad range of TAN loadings will ever prove more
accurate than fitting a linear model across the narrow
oligotrophic/mesotrophic range. Since the focus of this
paper is on mesotrophic/oligotrophic systems, the
parameter, t, was calculated for TAN concentrations
ranging between 0.1 and 0.5 g-TAN m
3. However, it
may be possible to extend the analysis beyond the range
that was analyzed in this study.
In summary, it is the author’s opinion that the linear
representation of data generated in the oligotrophic
and mesotrophic range will provide a more accurate
representation than the broadly calibrated Monod
kinetics. The development of a form of Monod
kinetics that is capable of representing heterotrophic
interference under eutrophic conditions is deemed
appropriate. Additional focus on data collection in the
oligotrophic zone is warranted. These data should be
used to confirm the assumption of zero intercept and to
verify the observations presented here.
Acknowledgements
This work was stimulated by a modeling effort
conducted under funding by Hubbs-SeaWorld
Research Institute of San Diego California. This
effort was under the direction of Mark Drawbridge.
Support for the analysis presented here was provided
by Bioclarifier Engineering Analysis and Design, LLC
of Baton Rouge Louisiana.
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