A Proposed Transient Model for
Cometabolism in Biofilm Systems
Pascale Champagne, Paul J. Van Geel, Wayne J. Parker
Department of Civil and Environmental Engineering, Carleton University,
Ottawa, Ontario, telephone: 613-520-2600, ext. 4628; fax: 613-520-3951;
e-mail: pchampag@ccs.carleton.ca
Received 11 November 1997; accepted 7 May 1998
Abstract: A dynamic model was developed to describe
the behaviour of primary and secondary substrates in
a biofilm reactor. The model incorporates structured ki-
netics to describe the generation and consumption of
reducing power in the catabolic and respiratory sub-
systems, respectively. Secondary substrate transforma-
tion through oxygenolytic or reductive mechanisms can
be modelled under either single or dual limitation of the
electron donor and electron acceptor substrates. An ex-
ample simulation of a theoretical biofilm system was
performed. © 1998 John Wiley & Sons, Inc. Biotechnol Bioeng
60: 541–550, 1998.
Keywords: biofilm; dual substrate limitation; cometabo-
lism; secondary substrate; biofilm modeling
INTRODUCTION
In biological wastewater treatment, many contaminants that
are of concern, if released into the environment, may be
subject to biodegradation through cometabolic processes
(e.g., quinoline (Malmstead et al., 1995), carbon tetrachlo-
ride (Bae and Rittmann, 1990)). Cometabolism refers to the
transformation of a secondary substrate by metabolic reac-
tions that do not contribute to the growth of microorgan-
isms. These reactions are significant in terms of secondary
degradation because the secondary substrates may be pre-
sent at concentrations below those required to sustain a
microbial population, or because the rate-limiting reactions
require energy that must be provided by a primary substrate.
Biodegradation of these compounds will hence be linked to
the presence and utilization of the primary substrates, which
consist of electron donors and electron acceptors. These
primary substrates influence the quantity of active biomass
that is present, and may either provide energy for cometa-
bolic reactions when metabolized, or act as a reactant in the
transformation reaction.
Numerous studies have demonstrated the relationship be-
tween primary substrate utilization and biomass growth un-
der single substrate limiting conditions (Annachhtre and
Khanna, 1987; Atkinson and Davies, 1974; Belhadir et al.,
1988; Capdeville et al., 1988; Hamoda, 1989; Kim and
Suidan, 1989; Lawrence and McCarty, 1970; Rittman,
1984; Rittmann and Manem, 1992; Rittmann and McCarty,
1978, 1980, 1981; Sa ´ez and Rittmann, 1988, 1990; Sa ´ez et
al., 1984; Shamat and Maier, 1980; Skowlund, 1990; Suidan
and Wang, 1985; Suidan et al., 1987; Tabak et al., 1990;
Trulear and Characklis, 1982; Wanner and Gujer, 1984). In
general, the Monod relationship has been found to ad-
equately represent this relationship. Fewer studies have ex-
amined growth when both the electron donor and electron
acceptor are limiting (Barton and McKeown, 1986; Borden
and Bedient, 1986; Howell and Atkinson, 1976; Kissel et
al., 1984; Lau et al., 1984; Odencrantz et al., 1990; Ritt-
mann and Dovantsiz, 1983; Sinclair and Ryder, 1975;
Sykes, 1973; Williamson and McCarty, 1976a,b). However,
recent studies (Bae, 1992; Bae and Rittmann, 1996a,b) have
indicated that a multiplicative double Monod relationship
adequately represents this behavior.
Several relationships between primary substrates and the
degradation of secondary substrates through oxygenolytic
and reductive reactions have been described in the litera-
ture: Hooker et al. (1994), Bae and Rittmann (1995), and
Wrenn and Rittmann (1995, 1996). The model of Hooker et
al. (1994) was derived through responses observed in ex-
perimental studies on the biodegradation of CCl
4
by deni-
trifying bacteria. The models developed by Wrenn and Ritt-
mann (1995, 1996) and Bae and Rittmann (1995) relate the
rate of transformation of a secondary substrate to the inter-
nal reducing power of the biomass, as indicated by the ratio
of reduced to oxidized nicotinamide adenine dinucleotide
(NADH/NAD) and the ratio of adenosine triphosphate to
adenosine diphosphate (ATP/ADP). These ratios are influ-
enced by the rate of metabolism of the electron donor and
the relative abundance of the electron acceptor.
In suspended growth systems, the bulk-phase concentra-
tions of the primary substrates provide the main driving
force for secondary substrate transformation. However, in
biofilm systems, mass transfer and diffusive limitations
within the biofilm will influence the relative concentrations
of primary and secondary substrates, as well as the level of
biomass activity. In addition, the concentrations of electron
donor and electron acceptor relative to each other will also
change through the biofilm depth. Thus, the rate of cometa-
bolic biodegradation of a secondary substrate can also be
expected to vary through the depth of the biofilm. Hence,
Correspondence to: Pascale Champagne
Contract grant sponsors: The Natural Sciences and Engineering Re-
search Council of Canada
© 1998 John Wiley & Sons, Inc. CCC 0006-3592/98/050541-10