Sunteți pe pagina 1din 4

Letters in Applied Microbiology ISSN 0266-8254

ORIGINAL ARTICLE

Re-interpretation of the logistic equation for batch


microbial growth in relation to Monod kinetics
F. Kargi
Department of Environmental Engineering, Dokuz Eylul University, Buca, Izmir, Turkey

Abstract

Keywords
batch microbial growth, kinetic constants,
logistic equation, Monod kinetics.
Correspondence
Fikret Kargi, Department of Environmental
Engineering, Dokuz Eylul University, Buca,
Izmir, Turkey. E-mail: fikret.kargi@deu.edu.tr

2008 1013: received 16 June 2008, revised


and accepted 31 October 2008
doi:10.1111/j.1472-765X.2008.02537.x

Aims: To determine the underlying substrate utilization mechanism in the


logistic equation for batch microbial growth by revealing the relationship
between the logistic and Monod kinetics. Also, to determine the logistic rate
constant in terms of Monod kinetic constants.
Methods and Results: The logistic equation used to describe batch microbial
growth was related to the Monod kinetics and found to be first-order in terms
of the substrate and biomass concentrations. The logistic equation constant
was also related to the Monod kinetic constants. Similarly, the substrate utilization kinetic equations were derived by using the logistic growth equation and
related to the Monod kinetics.
Conclusion: It is revaled that the logistic growth equation is a special form of
the Monod growth kinetics when substrate limitation is first-order with respect
to the substrate concentration. The logistic rate constant (k) is directly proportional to the maximum specific growth rate constant (lm) and initial substrate
concentration (S0) and also inversely related to the saturation constant (Ks).
Significance and Impact of the Study: The semi-empirical logistic equation can
be used instead of Monod kinetics at low substrate concentrations to describe
batch microbial growth using the relationship between the logistic rate constant
and the Monod kinetic constants.

Introduction
Microbial growth can simply be described by the following bioreaction
RS X ! nX RP

where S, X and P are the substrate, biomass and the


product respectively. The reaction is autocatalytic as the
biocatalyst (biomass) is generated by the reaction.
The rates of microbial growth, substrate utilization and
product formation have the following forms,

398

dX=dt f S; X; P

dS=dt gS; X; P 1=Y x=s dX=dt

dP=dt yS; X; P Y p=s dS=dt

Different equations were used to describe batch microbial growth (Shuler and Kargi 2002). Among the most
widely used growth rate expressions are the logistic equation and the Monod kinetics. The logistic equation developed by Verhulst considers only biomass concentration
disregarding the substrate utilization. Variation of biomass concentration with time is sigmoidal for the logistic
equation (Verhulst 1845). Monod kinetics considers both
the biomass and the rate limiting substrate concentrations in growth rate expression and is the most widely
used kinetic equation for batch microbial growth
(Monod 1949). However, the logistic equation has also
been used by many investigators to describe batch microbial growth (Simkins and Alexander 1984; Schimidt
et al.1985; Kingsland 2002). The kinetic constants in
those two equations were considered to be different and
not interrelated. While the Monod kinetics is mechanistic

2009 The Author


Journal compilation 2009 The Society for Applied Microbiology, Letters in Applied Microbiology 48 (2009) 398401

F. Kargi

Batch microbial growth kinetics

with meaningful constants, the logistic equation is a


mathematical approximation with a rather meaningless
rate constant.
Researchers investigated the suitability of both models
at different stages of microbial growth and found one
model or the other more suitable (Simkins and Alexander
1984; Schimidt et al. 1985). However, the relationship
between the two models has not been revealed theoretically to guide the researchers for the use of the right
model. Therefore, the major objective of this article is to
reveal the relationship between the logistic and Monod
kinetic equations for batch microbial growth and to
express the logistic rate constant in terms of Monod
kinetic constants.

The logistic equation used to describe the batch microbial


growth has the following form (Shuler and Kargi 2002;
Weisstein 2008).


dX
X
kX 1 
5
dt
Xm
where Xm is the maximum biomass concentration at the
end of batch growth and X is the biomass concentration
at any time during batch growth which are given by the
following equations.
X m X 0 Y x=s S0  Se

X X 0 Y x=s S0  S

6a

where X0 is the initial biomass concentration (gX l)1),


Yx s is the growth yield coefficient (gX g)1S), S0 and Se
are the substrate concentrations at the beginning (t = 0)
and at the end of batch growth (gS l)1) and S is the
substrate concentration at any time during batch growth
(gS l)1). X0 and Se values are usually negligible as
compared with Xm and S0.
With negligible X0 and Se, the eqns (6) and (6a) can be
approximated to
X m  Y x=s S0

6b

X Y x=s S0  S

6c

The integral form of the logistic growth equation has


the following form with the initial condition of X = X0 at
t = 0.
X0 ekt
1  X0 =X m 1  ekt

dX
lm S

X lX
dt
K s S

5a

Substrate concentration is not considered as a parameter


in the logistic growth equation. However, it is well known

where l and lm are the actual and the maximum specific


growth rates (h)1), S is the rate limiting substrate (nutrient) concentration (gS l)1), Ks is the saturation or halfrate constant (g l)1) and X is the biomass concentration
(gX l)1). Equation (6a) can be used to relate the biomass
and substrate concentrations during batch growth.
Substitution of
S S0 

Theoretical background

that the microbial growth rate is related to the rate-limiting substrate concentration according to the Monod
kinetics as follows (Monod 1949; Shuler and Kargi 2002).

X  X0
Y x=s

into eqn (7) yields the following growth rate equation.


lm Y x=s S0 X 0  X
dX

X
K s Y x=s Y x=s S0 X 0  X
dt

7a

Integration of eqn (7a) yields the following equation


describing the variation of the biomass concentration
with time during batch growth according to the Monod
kinetics (Shuler and Kargi 2002).
ALnX=X 0  BLnY x=s S0 X 0  X=Y x=s S0 lm t 7b
where
A = (KsYx s + S0Yx s + X0) (Yx sS0 + X0)
and
B = KsYx s (X0 + Yx sS0).
Both the differential and the integral forms of the
logistic and Monod kinetic equations are quite different. Monod kinetics is a mechanistic model where the
growth rate is related to both the substrate and
biomass concentrations as presented in eqn (7). The
logistic equation is independent of the substrate concentration and is only related to the biomass concentration. The growth rate is proportional to the actual
biomass concentration (X) and the difference between
the maximum and the actual biomass concentrations
(Xm ) X) which is known as the carrying capacity.
The logistic equation does not reveal any mechanism
for substrate utilization for batch growth unlike the
Monod kinetics.
The relationship between the logistic and Monod
kinetics for batch microbial growth
The logistic equation [eqn (5)] can be modified as follows
by using eqns (6) and (6a).

2009 The Author


Journal compilation 2009 The Society for Applied Microbiology, Letters in Applied Microbiology 48 (2009) 398401

399

Batch microbial growth kinetics

F. Kargi



dX
X
kX 1 
kX 0 Y x=s S0  S
dt
Xm


X 0 Y x=s S0  S
1
X 0 Y x=s S0

5b

eqn (8a) into eqn (7c) yields eqn (5) which is the differential form of the logistic equation.
Equation (7c) can be further arranged as follows by
neglecting X0

As X0 > X, X0 can be neglected and eqn (5b) reduces to


eqn (5c),
5c

Equation (5c) indicates the inherent first-order kinetics


assumption in the logistic equation as the eqn (5c) is
first-order in terms of both the substrate and the biomass
concentrations.
By using the following relationship,
dX
dS
Y x=s
dt
dt

eqn (5c) can be written in terms of the substrate concentration as follows:




dS
S
S
 kS0  S kS 1 
9
dt
S0
S0
Equation (9) is the logistic equation for substrate utilization in batch growth. Variation of the substrate concentration with time during batch growth can be estimated
by the following equation,
S S0 

X  X0
Y x=s

dX
l S
lX m X
dt
Ks

7c

dS dX 1
l S

m S0  S
dt
dt Y x=s
Ks

7d

lm
Ks

Sk0 , then eqn (7d) takes the following form


d
 S kS1  S=S0 which has the same form as eqn
dt
(9).
Therefore, the Monod kinetics reduces to the logistic
equation with the assumption of first-order kinetics with
respect to the rate limiting substrate. In other words, the
logistic equation is a special form of Monod kinetics for
batch microbial growth with the assumption of the firstorder kinetics which holds for low substrate concentrations (S > Ks).
The batch growth and substrate utilization curves as
estimated by the Monod [eqn (7b)], logistic [eqn (5a)]
and the first-order Monod [eqn (7c)] equations were
compared in Fig. 1 by using the following constants.

As

X 0 05 g l1 ;
K s 2 g l1 ;

8a

where X = X0 and S = S0 at t = 0 yielding X = Xm and


S = Se at the end of batch growth. Biomass concentration (X) in eqn (8a) is to be estimated by using
eqn (5a).
The Monod kinetics for batch microbial growth as represented by eqn (7) reduces to the first-order kinetics at
low substrate concentrations (S > Ks) as follows:

7c

or in terms of substrate utilization rate

S0 10 g l1 ;

lm 025 h1 ;

Y x=s 035 gX g1 S

The first-order Monod and the logistic equations


yielded the same curves for the variation of biomass and
12
10
X, S (g l1)

dX
S
S
kY x=s S0  S kX
dt
S0
S0

dX
l S
l S
lX m X m Y x=s S0  S
dt
Ks
Ks

8
6
X

by comparing eqns (5c) and (7c) one can easily find out
that the rate constant for the logistic equation is related
to the Monod kinetic constants as follows:

l
k m S0 k 1 S0
Ks

2
S

10

where k1 (lm Ks) is the first-order rate constant for the


Monod kinetics. The rate constant for the logistic equation is directly related to the first-order rate constant and
the initial substrate concentration (S0). Substitution of
400

4
Time (h)

Figure 1 Batch microbial growth and substrate utilization curves as


predicted by the logistic (
), Monod (
) and the first-order Monod
() kinetics. X0 = 05 g l)1, S0 = 10 g l)1, lm = 025 h)1, Ks = 2 g
l)1, Yx s = 035 gX g)1S.

2009 The Author


Journal compilation 2009 The Society for Applied Microbiology, Letters in Applied Microbiology 48 (2009) 398401

F. Kargi

the substrate with time. However, the growth and substrate consumption curves estimated by the Monod equation were quite different and sluggish as compared with
the logistic curve due to saturation type substrate consumption in Monod kinetics.
Conclusion
The semi-empirical logistic batch growth equation was
related to the mechanistic Monod kinetics and the mechanism of substrate utilization was revealed for the logistic
growth curve. The logistic equation was found to be a
special case of the Monod kinetics with the first-order
substrate utilization kinetics. The Monod kinetics reduces
to the logistic equation when the growth kinetics is firstorder in terms of the substrate and the biomass concentrations. The logistic rate constant (k) was found to be
proportional to the maximum specific growth rate constant (lm) and the initial substrate concentration (S0) and
also inversely proportional to the saturation constant
(Ks). The logistic growth equation can be used instead of
complicated Monod kinetics to describe batch microbial
growth at low substrate concentrations where the rate of
substrate utilization is first-order with respect to substrate
concentration.

Batch microbial growth kinetics

References
Kingsland, S.E. (2002) Modeling Nature. Chicago, IL, USA:
The University of Chicago Press.
Monod, J. (1949) The growth of bacterial cultures. Annu Rev
Microbiol 8, 371374.
Schimidt, S.K., Simkins, S. and Alexander, M. (1985) Models
for the kinetics of biodegradation of organic compounds
not supporting growth. Appl Environ Microbiol 50, 323
331.
Shuler, M.L. and Kargi, F. (2002) Bioprocess Engineering: Basic
Concepts, 2nd edn. NJ, USA: Prentice Hall.
Simkins, S. and Alexander, M. (1984) Models for mineralization kinetics with the variables of substrate concentration
and population density. Appl Environ Microbiol 47, 1299
1306.
Verhulst, P.F. (1845) Recherches mathematiques sur la loi
daccroissement de la population. LAcademie Royale des
Sci. Et Belles-Lettres de Bruxelles 18, 141.
Weisstein, E.W. (2008) Logistic equation. MathWorldA Wolfram Web Resource. http://mathworld.wolfram.com/Logistic
equation.html.

2009 The Author


Journal compilation 2009 The Society for Applied Microbiology, Letters in Applied Microbiology 48 (2009) 398401

401

S-ar putea să vă placă și