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Abstract
Nitrification is classified as a two-step consecutive reaction where R1 represents the rate of formation of the intermediate
product NO2-N and R2 represents the rate of formation of the final product NO3-N. The relative rates of R1 and R2 are
influenced by reactor type characterized hydraulically as plug-flow, plug-flow with dispersion and mixed-flow. We develop
substrate conversion models for fixed-film biofilters operating in the first-order kinetic regime based on application of
chemical reactor theory. Reactor type, inlet conditions and the biofilm kinetic constants Ki (h1) are used to predict changes
in NH4-N, NO2-N, NO3-N and BOD5. The inhibiting effects of the latter on R1 and R2 were established based on the g
relation, e.g.:
N
Ki ½BOD5
g¼ ¼ 1
Ki;max ½BOD5;max
where BOD5,max is the concentration that causes nitrification to cease and N is a variable relating Ki to increasing BOD5.
Conversion models were incorporated in spreadsheet programs that provided steady-state concentrations of nitrogen and
BOD5 at several points in a recirculating aquaculture system operating with input values for fish feed rate, reactor volume,
microscreen performance, make-up and recirculating flow rates. When rate constants are standardized, spreadsheet use
demonstrates plug-flow reactors provide higher rates of R1 and R2 than mixed-flow reactors thereby reducing volume
requirements for target concentrations of NH4-N and NO2-N. The benefit provided by the plug-flow reactor varies with
hydraulic residence time t as well as the effective vessel dispersion number, D/mL. Both reactor types are capable of
providing net increases in NO2-N during treatment but the rate of decrease in the mixed-flow case falls well behind that
predicted for plug-flow operation. We show the potential for a positive net change in NO2-N increases with decreases in the
dimensionless ratios K2;ðR2 Þ =K1;ðR1 Þ and [NO2-N]/[NH4-N] and when the product K1;ðR1 Þ t provides low to moderate NH4-N
conversions. Maintaining high levels of the latter reduces the effective reactor utilization rate (%) defined here as (RNavg/
RNmax)100 where RNavg is the mean reactive nitrogen concentration ([NH4-N] + [NO2-N]) within the reactor, and RNmax
represents the feed concentration of the same. Low utilization rates provide a hedge against unexpected increases in
substrate loading and reduce water pumping requirements but force use of elevated reactor volumes. Further g effects on R1
* Corresponding author. Tel.: +1 304 724 4425; fax: +1 304 724 4428.
E-mail addresses: barnaby_watten@usgs.gov (B.J. Watten), philip_sibrell@usgs.gov (P.L. Sibrell).
and R2 can be reduced through use of a tanks-in-series versus a single mixed-flow reactor configuration and by improving the
solids removal efficiency of microscreen treatment.
Published by Elsevier B.V.
performance as measured by required volume, T = 24 8C). When S is much greater than Ks, Equation
pumping requirements and changes in water chemistry (3) reduces to an expression that is zero-order with
(Rittmann, 1982; Levenspiel, 1972; Weber, 1972). We respect to S. When S is limiting and smaller than Ks, as
quantify these effects following development of in typical salmonid aquaculture applications Equation
performance algorithms that allow standardization (3) reduces to an expression that is first-order with
of certain steady-state biofilm kinetic parameters. respect to S. Here, the rate expressions for each
Model use focuses on both organic effects and nitrogen component of the reaction can be expressed
incomplete nitrification given the sensitivity of fish as (Levenspiel, 1972; Srna and Baggaley, 1975;
to NO2-N at relatively low concentrations. Weatherley, 1984):
1 dMNH4 -N
rNH4 -N ¼ ¼ K1 ½NH4 -N (4)
2. Model development V dt
1 dMNO2 -N
2.1. Nitrification rNO2 -N ¼ ¼ K1 ½NH4 -NK2 ½NO2 -N
V dt
(5)
Nitrification within the biofilm is classified as a
1 dMNO3 -N
two-step consecutive reaction where R1 represents the rNO3 -N ¼ ¼ K2 ½NO2 -N (6)
rate of formation of the intermediate product NO2-N V dt
and R2 represents the rate of formation of the final where M is the mass substrate reacted, V reactor
product NO3-N. The stoichiometry of the reaction has volume and K1 and K2 are the empirically derived
been described as (Haug and McCarty, 1972): first-order reaction rate constants (time1) that repre-
sent the product of a substrate utilization rate constant
R1 : 55NH4 þ þ 5CO2 þ 76O2 and the active microbial mass per unit volume of the
! C6 H7 O2 N þ 54NO2 þ 52H2 O þ 109Hþ reactor. Chemical reaction kinetics, as described in the
(1) equations above, have been used with success to
follow the progress of nitrification within fixed-film
R2 : 400NO2 þ 5CO2 þNH4 þ þ 195O2 þ2H2 O reactors (O’Shaughnessy et al., 1984; Srna and Bag-
! C6 H7 O2 N þ 400NO3 þ Hþ (2) galey, 1975; Weatherley, 1982, 1984; Weatherley
et al., 1993; Watten et al., 1993, 1998). Values of
where C6H7O2N represents the yield (mass) of bac- K1 and the ratio K2/K1 will vary with reactor type and
terial cells. The reactions above are further classified application. Srna and Baggaley (1975) established rate
as being heterogeneous, i.e., fluid and solid phases are constants for a marine subgravel filter system operat-
present and necessary for the reactions to proceed. In ing with an inlet NH4-N of 0.1–0.7 mg/L. Tempera-
both cases, reaction kinetics follow the mixed-order ture, pH and sulfide ion (0.03 mg/L) affected K1 and
Monod relationship (Benefield and Randall, 1980): K2 but small changes in salinity did not (5–8 ppt). K1
dS km xS (h1) rose with increasing pH whereas K2 (h1) fell as
¼ (3) described in the relations: K1 = 0.230 (pH) 1.6
dt Ks þ S
(7.4 < pH < 7.8); K2 = 0.053 (pH) +0.531
where (dS/dt) is the overall substrate utilization rate, (6.6 < pH < 8.6). O’Shaughnessy et al. (1984) pre-
mass volume1 time1, km the maximum rate of sub- sent rate constants for a horizontal plug-flow reactor
strate utilization per unit of attached biomass, time1, 25 m in length. K1 over the NH4-N range of 1.0–
S the substrate concentration, mass volume1, Ks the 1.5 mg/L averaged just 0.039 h1 reflecting the low
saturation constant which has a value equal to the specific surface area provided by the reactor design
substrate concentration when (dS/dt)/x = 0.5km, and x (156 m2/m3). K2/K1 averaged 1.91. Weatherley (1984)
is the active biomass concentration, mass volume1. correlated K1 and K2 with pH based on Srna and
Typical values for Ks;NH4 -N and Ks;NO2 -N are, respec- Baggaley (1975), temperature as described by an
tively, 0.5–2.0 mg/L (Benefield and Randall, 1980) Arrhenius type relationship and dissolved oxygen
and 2.7 mg/L (Rittmann and Snoeyink, 1984; (DO) following the observations of Knowles et al.
B.J. Watten, P.L. Sibrell / Aquacultural Engineering 34 (2006) 198–213 201
(1965). A rate modifying factor for the latter was used where t is expressed in hours. Nitrification is, for all
to describe a linear reduction in K1 with decreases in practical purposes, a unimolecular type reaction
DO within the threshold range 0.7–2.0 mg/L and for (Equation (1) and (2)). Therefore, the following nitro-
K2 within the threshold range 0.7–4.0 mg/L. Predicted gen balance holds:
Ki values agreed well with values established for a
subgravel filter during model verification runs. Here, ½NH4 Nin ¼ ½NH4 -Nout þ ½NO2 -Nout þ ½NO3 -Nout
K1 and K2 averaged 0.430 and 0.100 h1, respectively, (9)
providing a K2/K1 ratio of 0.233. Ebeling (2000)
correlated K1 with hydraulic loading (HL) during an which leads to a solution for [NO3-N]out (Weber,
evaluation of a commercial bubble-wash bead filter 1972):
(T = 21–23 8C). K1 increased with HL from a low of
12.0 h1 at HL = 13.2 L/min to a high of 35.5 h1 at K1 eK2 t K2 eK1 t
½NO3 -Nout ¼ ½NH4 -Nin 1 þ
HL = 80.6 L/min. Brazil (2005) reported rate con- K2 K1 K2 K1
stants for a commercial scale RBC operating at a (10)
temperature of 28 8C. In this case, K1 averaged
2.4 h1 with a K2/K1 average of 4.5. Watten et al. Equations (8)–(10) assume NO2-N and NO3-N are
(1993) reported a mean K1 and K2/K1 ratio of 13.9 h1, not present in the influent, an unusual case in salmonid
and 1.6 for a fluidized bed of plastic beads (960 m2/ aquaculture operations. Correcting Equations (8)–(10)
m3) operating with an inlet NH4-N of 0.2–1.5 mg/L for inlet concentrations above zero results in:
(14.3 8C). Concurrently, K1 and the ratio K2/K1 varied
with location in a second reactor treatment of equal ½NO2 -Nout
volume that consisted of three fluidized bed compo- K1 ½NH4 -Nin K1 t K2 t
nents operated in series—K1 and the ratio K2/K1 were ¼ ðe e Þ þ ½NO2 -Nin eK2 t
K2 K1
15.8 h1 and 2.8 for stage 1, 20.3 h1 and 1.5 for stage (11)
2, and 12.8 h1 and 3.1 for stage 3. Staging effects on
Ki values have also been reported for RBC applica-
tions (Brazil, 2005) reflecting changes in component K1 eK2 t K2 eK1 t
½NO3 -Nout ¼ ½NH4 -Nin 1 þ
feed composition and substrate loading. Clearly, Ki K2 K1 K2 K1
values vary widely and must therefore be established þ ½NO2 -Nin ð1 eK2 t Þ þ ½NO3 -Nin
for a specific set of operating conditions either empiri- (12)
cally or through application of theoretical models like
those of Rittmann (1982) and Rittmann and McCarty The rate constant K1 is calculated based on
(1980, 1981). Equation (7) given NH4-N conversion, t and the
relation:
2.2. Plug-flow conversion
½NH4 -Nout
Integration of Equations (4) and (5) gives effluent K1 t ¼ ln (13)
½NH4 -Nin
NH4-N, NO2-N and NO3-N concentrations versus
reactor retention time t when the influent NH4-N The rate constant K2 is solved for by trial and error
concentration is known. In the plug-flow reactor case, given [NH4-N]in, [NO2-N]in, [NO2-N]out, t, Equation
we have (Levenspiel, 1972; Srna and Baggaley, 1975; (11), and a corresponding K1 from Equation (13).
Weatherley, 1982, 1984):
2.3. Mixed-flow conversion
½NH4 -Nout ¼ ½NH4 -Nin eK1 t (7)
Integration of the rate expressions rNH3 -N (Equation
(4)) and rNO2 -N (Equation (5)) for the mixed-flow
K1 ½NH4 -Nin K1 t reactor case gives alternate conversion models for each
½NO2 -Nout ¼ ðe eK2 t Þ (8)
K2 K1 of the nitrogen components of interest (Levenspiel,
202 B.J. Watten, P.L. Sibrell / Aquacultural Engineering 34 (2006) 198–213
K2 is solved for by trial and error given [NH4-N]in, The rate constants K1 and K2 used to establish
[NO2-N]in, [NO2-N]out, t, Equation (17) and a corre- reactor performance will vary with operating condi-
sponding K1 from Equation (19). tions including the feed concentration of degradable
organic matter that stimulates heterotrophic bacterial
2.4. Tanks-in-series conversion growth (Bovendeur et al., 1990; Ohashi et al., 1995;
Zhu and Chen, 2001). The stoichiometry of the
The conversion models presented thus far assume oxidation reaction associated with organic matter has
ideal plug-flow or mixed-flow reactor behavior. Ideal been expressed as (Benefield and Randall, 1980):
reactor behavior is difficult to achieve, particularly in C5 H12 þ 8O2 ! 5CO2 þ 6H2 O (23)
the plug-flow case, given the sensitivity of RTD to
system geometry. The vessel dispersion coefficient, D/ Although reaction kinetics adhere to the mixed-
mL, is used to identify reactor type given the measured order Monod relationship (Equation (3)), S in
mean residence time, t, and variance, s2, of a salmonid RAS applications (e.g., BOD5) is typically
conservative tracer. These variables are related with much smaller than Ks resulting in kinetics that are
B.J. Watten, P.L. Sibrell / Aquacultural Engineering 34 (2006) 198–213 203
Fig. 2. Spreadsheet model components used to simulate the effect of reactor type on water quality dynamics in an RAS. The letters (a–j)
represent model prediction points for the variables BOD5, NH4-N, NO2-N and NO3-N.
Model inputs included reactor type (plug-flow or addressed here but was in a similar spreadsheet
mixed-flow), K1, the ratio K2/K1, system make-up flow designed to simulate the effects of a microscreen
rate (Qmf, m3/h), recirculating flow rate (Qrf, m3/h), BOD5 removal step and reactor staging. Major com-
make-up flow composition ([NH4-N]mf, [NO2-N]mf, ponents of the system modeled are given in Fig. 2. Net
[NO3-N]mf, mg/L), reactor volume (V, m3) and fish BOD5 input was taken as the product of fish feed
feed rate (kg/d). The latter was used to calculate NH4- rate (kg/d), 1 E, and the ratio g biosolids/kg feed,
N loading based on the ratio 29 g/kg feed which where E represents the biosolids removal efficiency
represents a typical value for salmonids (Speece, of the microscreen as a decimal fraction. We used a
1973; Liao and Mayo, 1974). Net changes in nitrogen net biosolids (BOD5) production rate of 400 g/kg
components were established using the reactor feed based on the observations of Speece (1973)
specific conversion models described earlier (either and Liao and Mayo (1974). BOD5 was tracked through
Equations (7)–(12) or (14)–(18)) in combination with each RAS component with conversion accounted for
a material balance. For example, in the case of NH4-N in the primary mixed-flow reactor and tanks-in-series
the following balance holds: reactors (Equation (25)). Reactor specific BOD5
concentrations were used to correct K1 and K2 for
Qmf ½NH4 -Nmf þ MNH4 N ¼ Qof ½NH4 -Nof inhibition effects given a selected value for N
þ ðQmf þ Qrf Þð½NH4 -Nin ½NH4 -Nout Þ (27) (Equation (26)). Additional program inputs included
reactor volumes and reactor configuration—either a
where in this case NH4-N concentrations are single mixed-flow reactor, a tanks-in-series reactor
expressed as g/m3, Qof is a system overflow rate (six stages) or a combination of the both types as
(Qof = Qmf, m3/h) and MNH4 -N is the mass of NH4- illustrated in Fig. 2. In each of the three spreadsheet
N produced by the fish (g/h). The spreadsheet was programs we assume dissolved oxygen, CO2, tem-
designed to allow system operation without recircula- perature, pH and alkalinity are maintained within
tion, if desired, so as to simulate single pass or post desirable ranges by RAS system components not
treatment applications of the biofilm reactors, i.e., included in our modeling exercise and that the unit
Qrf = 0. When recirculation is employed (Qrf > 0), processes/operations used here do not influence the
an iteration procedure and check for convergence reactants of interest. Further, and following the RAS
was required given the rise in reactant concentrations models of Weatherley (1982, 1984) and Weatherley
that occur with time when reactant conversion per pass et al. (1993), we do not address conversion of BOD5,
is less than 100%. Organic inhibition was not NH4-N or NO2-N that may occur outside of the
B.J. Watten, P.L. Sibrell / Aquacultural Engineering 34 (2006) 198–213 205
3. Model application
Fig. 9. Effect of K1t and the ratio [NO2-N]in/[NH4-N]in on the ratio [NO2-N]out/[NO2-N]in when operating a plug-flow reactor.
both reactor types, the high NH4-N conversion rates potential (g N/(m3 d)). For example, line A, Fig. 3,
required to establish the condition [NO2-N]out/[NO2- gives the sum of NH4-N and NO2-N versus location
N]in 1.0 lead to a substantial degree of under- along the longitudinal axis of a plug-flow reactor
utilization of the biofilm substrate conversion constructed with a uniform cross-sectional area. We
define the sum of these two nitrogen components as
reactive nitrogen (RN) given their combined or
individual abilities to sustain nitrification reactions.
Here, the microbes within the reactor work at a rate
that is fixed, in the first-order regime, by a mean RN
concentration that lies well below inlet levels
(RNmax). The effective RN concentration (RNavg)
can be calculated by applying the mean value
theorem of integral calculus given functions describ-
ing either the individual or combined (RN) nitrogen
component curves. Considering the combined func-
tion, we know from our material balance (Equation
(3)) that at any time t:
solid phase, e.g., a fluidized bed. The K3 value required gives g and BOD5 versus position as well as model
for BOD5 conversion estimates (6.3 h1; Equation predicted NH4-N, NO2-N and NO3-N when operating
(25)) was taken from Watten et al. (1993). The product- at feed rates of 2 and 12 kg/d. Note g increases, at
poisoning coefficient N (Equation (26)) was fixed at a 12 kg/d, from a low of 0.13 in the primary reactor to a
value of 5 to explore a case of high initial sensitivity to high of 0.61 in the last stage of the series reactor. An
BOD5 (Fig. 1). BODmax (Equation (26)) was set at initial rise then fall in NO2-N can be seen as water
20 mg/L, for example purposes only, based on the moves from stage to stage following the response
observations of Antonie (1976) and Weng and Molof predicted in Fig. 7. Further runs (Table 1) show
(1974). The make-up flow requirement of the RAS in all steady-state concentrations of NO2-N and NH4-N
cases (0.1 m3/h) represented 1% of the total flow drop significantly when the poisoning coefficient N is
directed through the rearing unit (10 m3/h). The reduced from 5 to 1, the latter establishing a linear
removal of suspended solids across the microscreen (negative) effect of BOD5 on g (Fig. 1), e.g., in the fish
was initially held at 60% (E = 0.6). This lies within the tank effluent NO2-N and NH4-N drop from 0.55
range of E = 0.21 0.68 reported for RAS applications to 0.14 and 1.99 to 1.55 mg/L, respectively. The g
by Bergheim and Brinker (2003) and Twarowska et al. factor here rises to a range of 0.61–0.91. Reducing
(1997). Fish feed rates were varied from 2 to 12 kg/d the microscreen efficiency to 30% (E = 0.3) acts to
which correspond to a change in dissolved oxygen lower g and elevate BOD5, NO2-N and NH4-N
across the rearing unit of between 2.1 and 12.5 mg/L (Table 1). Fig. 13 shows this effect on fish tank
assuming an oxygen demand ratio of 0.25 kg/kg feed effluent composition over the solids removal effi-
(Losordo and Westers, 1994). Model predicted con- ciency range of 10–80%. In this case the N used to
centrations of BOD5 in the microscreen effluent define g was fixed at 3. Note that NH4-N and NO2-N
(reactor feed) ranged from 2.0 to 12.2 mg/L in the drop at a decreasing rate as solids removal efficiency
mixed-flow case and from 1.6 to 9.9 mg/L in the tanks- approaches 80%. The E-value required for a design
in-series case. The latter provided lower NH4-N and level of NH4-N and NO2-N are lower in the tanks-in-
NO2-N concentrations than the mixed-flow reactor series versus mixed-flow reactor configuration.
(Fig. 12) in agreement with our earlier simulation
results established without BOD5 inhibition (Figs. 4–6).
The inhibition parameter g (Equation (26)), in the 4. Summary
mixed-flow case, dropped from 0.92 to 0.31 as feed rate
increased. In the tanks-in-series application, g varied We develop substrate conversion models for fixed-
with position along the length of the reactor axis film biological filters operating in the first-order
reflecting stage-by-stage reductions in BOD5. Table 1 kinetic regime based on application of chemical
Table 1
Model predicted water chemistry variables vs. sample point in an RAS operating under four test case conditions
Test case Variable RAS component
211
212 B.J. Watten, P.L. Sibrell / Aquacultural Engineering 34 (2006) 198–213
Science, vol. 27. Elsevier Science, Amsterdam, The Nether- Shnel, N., Barak, Y., Ezer, T., Dafni, Z., Van Rijn, J., 2002. Design
lands, pp. 9–60. and performance of a zero discharge tilapia recirculating system.
Manthe, D.P., Malone, R.F., Kumar, S., 1984. Limiting factors Aquacult. Eng. 26, 191–203.
associated with nitrification in closed blue crab shedding sys- Smith, C.E., Williams, W.G., 1974. Experimental nitrate toxicity in
tems. Aquacult. Eng. 3, 119–140. rainbow trout and Chinook salmon. Trans. Am. Fish. Soc. 103,
Meunier, A.D., Williamson, K.J., 1981. Packed bed biofilm reactors: 389–390.
simplified model. J. Environ. Eng. Division, ASCE EE2, 307– Speece, R.E., 1973. Trout metabolism characteristics and the
315. rational design of nitrification facilities for water reuse in
Miller, G.E., Libey, G.S., 1984. Evaluation of a trickling biofilter in a hatcheries. Trans. Am. Fish. Soc. 2, 323–334.
recirculating aquaculture system containing channel catfish. Srna, R.F., 1975. A modular nitrification filter design based on a
Aquacult. Eng. 3, 39–57. study of the kinetics of nitrification of marine bacteria. Proc.
Nam, T.K., Timmons, M.B., Montemagno, C.D., Tsukuda, S.M., World Maricult. Soc. 6, 463–478.
2000. Biofilm characteristics as affected by sand-size and loca- Srna, R.F., Baggaley, A., 1975. Kinetic response of perturbed marine
tion in fluidized bed vessels. Aquacult. Eng. 22, 213–224. nitrification systems. J. Water Polut. Control Fed. 47, 472–486.
Nijhof, M., 1995. Bacterial stratification and hydraulic loading Summerfelt, S.T., Cleasby, J.L., 1993. Hydraulics in fluidized-bed
effects in a plug-flow model for nitrifying trickling filters applied biological reactors. In: Wang, J.K. (Ed.), Techniques for Modern
in recirculating fish culture systems. Aquaculture 134, 49–64. Aquaculture. American Society of Agricultural Engineers, Saint
Nijhof, M., Klapwijk, A., 1995. Diffusional transport mechanisms Joseph, Michigan, pp. 439–457.
and biofilm nitrification characteristics influencing nitrite levels Twarowska, J.G., Westerman, P.W., Losordo, T.M., 1997. Water
in nitrifying trickling filter effluents. Water Res. 29, 2287–2292. treatment and waste characterization evaluation of an intensive
O’Shaughnessy, J.C., Blanc, F.C., Barker, T.G., 1984. Fixed film recirculating fish production system. Aquacult. Eng. 16, 133–147.
nitrification kinetics. In: Second International Conference on Tucker, C.S., 1985. Channel Catfish Culture. Elsevier Science
Fixed Film Processes, Pittsburgh, Pennsylvania, pp. 1389–1403. Publishers, Amsterdam, The Netherlands.
Ohashi, A., Viraj de Silva, D.G., Mobarry, B., Manem, J.A., Stahl, Watten, B.J., Timmons, M.B., Summerfelt, S.T., Vinci, B.J., 1998.
D.A., Rittmann, B.E., 1995. Influence of substrate C/N ratio on Comparative performance of biofilm reactor types: application
the structure of multi-species biofilms consisting of nitrifers and of steady-state biofilm kinetics. In: Libey, G.S., Timmons, M.B.
heterotrophs. Water Sci. Tech. 32, 75–84. (Eds.), Proceedings of the Second International Conference on
Paller, M.H., 1992. An analytical model for predicting the carrying Recirculating Aquaculture, Roanoke, Virginia, pp. 355–358.
capacity of submerged biofilters used in aquaculture. J. Appl. Watten, B.J., Ridge, W.J., Timmons, M.B., 1993. Evaluation of a
Aquacult. 1, 1–25. Multi-stage Fluidized Bed Biological Filter for Accelerated
Ridha, M.T., Cruz, E.M., 2001. Effect of biofilter media on water Nitrification and Reduced Energy Demand. American Fisheries
quality and biological performance of the Nile tilapia Oreo- Society Annual Meeting, Pittsburgh, Pennsylvania.
chromis niloticus L. reared in a simple recirculating system. Weatherley, L.R., 1982. Application of simple dynamic response
Aquacult. Eng. 24, 157–166. analysis to a recirculating aquaculture system—a preview.
Rittmann, B.E., 1982. Comparative performance of biofilm reactor Aquacult. Eng. 1, 93–113.
types. Biotechnol. Bioeng. 24, 1341–1370. Weatherley, L.R., 1984. Rate models for a marine biological filter.
Rittmann, B.E., McCarty, P.L., 1980. Evaluation of steady-state Aquacult. Eng. 3, 15–29.
biofilm kinetics. Biotechnol. Bioeng. 22, 2359–2373. Weatherley, L.R., Hill, R.G., Macmillan, K.S., 1993. Process mod-
Rittmann, B.E., McCarty, P.L., 1981. Substrate flux into biofilms of eling of an intensive aquaculture system. Aquacult. Eng. 12,
any thickness. Environ. Sci. Technol. 15, 831–849. 215–230.
Rittmann, B.E., Snoeyink, V.L., 1984. Achieving biologically Weber, W.J., 1972. Physiochemical Processes for Water Quality
stable drinking water. J. Am. Wastewater Assoc., October Control. John Wiley and Sons Inc., New York.
106–114. Weng, C., Molof, A.H., 1974. Nitrification in the biological fixed
Rogers, G.L., Klemetson, S.L., 1985. Ammonia removal in selected film rotating disk system. J. Water Pollut. Control. Fed. 46,
aquaculture water reuse biofilters. Aquacult. Eng. 4, 135–154. 1674–1685.
Rusten, B., McCoy, M., Proctor, R., Siljudalen, J.G., 1998. The Wheaton, F.W., Hockheimber, J.N., Kaiser, G.E., Krones, M.J.,
innovative moving bed biofilm reactor/solids contact reaeration Libey, G.S., Easter, C.C., 1994. Nitrification filter principles.
process for secondary treatment of municipal wastewater. Water In: Timmons, M.B., Losordo, T.M. (Eds.), Aquaculture Water
Environ. Res. 70, 1083–1089. Reuse Systems: Engineering Design and Management. Devel-
Sandu, S.I., Boardman, G.D., Watten, B.J., Brazil, B.L., 2002. opments in Aquaculture and Fisheries Science, vol. 27. Elsevier
Factors influencing the nitrification efficiency of fluidized bed Science, Amsterdam, The Netherlands, pp. 127–171.
filter with a plastic medium. Aquacult. Eng. 26, 41–60. Zhu, S., Chen, S., 1999. An experimental study on nitrification
Satoh, H., Okabe, S., Norimatsu, N., Watanabe, Y., 2000. Signifi- biofilm performance using a series reactor system. Aquacult.
cance of substrate C/N ratio on structure and activity of nitrify- Eng. 20, 245–259.
ing biofilms determined by in situ hybridization and use of Zhu, S., Chen, S., 2001. Effects of organic carbon on nitrification
microelectrodes. Water Sci. Tech. 41, 317–321. rate in fixed film biofilters. Aquacult. Eng. 25, 1–11.