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Kinetics is a study that deals with rate of the reaction. It is based on the

mechanisms of any processes-physical, chemical or biological. The chemical kinetics

deals with how the rate of reaction is dependent upon the concentration of the reactants.

In a biochemical reaction, kinetics is a little complicated. A living cell, during

fermentation or any biological processes, require essential nutrients and suitable

environment to survive. Hence a biochemical reaction proceeds with the intervention of

microbial cells in a liquid medium containing nutritive sources under conditions like pH,

temperature. The cells multiply and grow, consuming a part of substrate and other

necessary components from the fermentation broth. Cell growth is associated with two

Certain parameters/ phenomena that determine cell population kinetics are the

characteristics of culture broth, nutrients and substrates used for growth and production

of metabolites, cell to cell heterogeneity, microbial cells of different ages manifesting

metabolic activities and characterization of biochemical pathway (Figure 5.1). These

complex parameters make it difficult to formulate a simple kinetic model for cellular

activities (Bailley and Ollis, 1986). A kinetic model describes the behavior of the cellular

processes through possible mathematical equations and it serves to be a very effective

tool to test and eliminate the extremities. Different kinetic models have been proposed in

several research works for microbial growth, substrate utilisation and product formation

in various fermentation modes.

representations of cellular population are taken into consideration. An individual cell in a

liquid broth is a complicated multi component system, which is not spatially homogenous

even at a single cell level. Nutrient solution used for the inoculum growth and production

is also a multi component system containing all the complex nutrients for cell growth and

accumulates various products as the result of metabolic activities. According to number

of components used in microbial representations and considering cells as heterogenous

mass or wholly as average cell clusters, different perspective are put forward for cell

population kinetic representations (Figure 5.2).

determine cell population kinetics (Adapted from Bailley and Ollis,1986)

Structured model is one perspective in which the microbial cells are considered as

multicomponent systems. When cell population is treated as one component system, it is

referred to as unstructured model. The rate of reaction depends only on the parameters

inside the fermenter. These models only contain growth, substrate consumption and

product formation kinetics. Cell to cell heterogeneity and discrete cells compose a multi

component entity, constituting ultimately a segregated perspective. An unsegregated view

considers average cellular properties. In actual cellular cases, the situation is a structured

and segregated one. In balanced growth of cells, the metabolic activities are coordinated

in such a way that the average cell composition is not influenced by increase in cell mass.

In such cases, models that do not consider the multi component nature of cells are

necessary. Hence unsegregated, unstructured model is the most idealized situation when

analyzing the growth of cells (Bailley and Ollis, 1986).

Figure 5.2 Different perspectives for cell population kinetic representation (Adapted

from Bailley and Ollis,1986)

model, which considers substrate conversion to cell mass, to product and substrate

consumption maintenance,

ୢୗ ଵ ୢ୶ ଵ ୢ

ൌ െ െ ୶

ୢ୲ ଢ଼୶Ȁୱ ୢ୲ ଢ଼୮Ȁୱ ୢ୲

!

where Yx/s is the yield coefficient for biomass with respect to substrate consumed and Yp/s

is the yield coefficient for product formed with respect to the substrate consumption.

ୟୱୱ୭ୡୣ୪୪୭୰୫ୣୢ ୶ ୶ି୶బ

୶Ȁୱ ൌ ൌ ൌ

ୟୱୱ୭ୱ୳ୠୱ୲୰ୟ୲ୣୡ୭୬ୱ୳୫ୣୢ ୱ ୱబ ିୱ

" !

ୟୱୱ୭୮୰୭ୢ୳ୡ୲୭୰୫ୣୢ ିబ

Ȁୱ ൌ ൌ ൌ

ୟୱୱ୭ୱ୳ୠୱ୲୰ୟ୲ୣୡ୭୬ୱ୳୫ୣୢ ୱ ୱబ ିୱ

# !

If the amount of carbon sources used for product formation and maintenance

constants are neglected when modeling substrate consumption rate, the model becomes,

ୢୗ ଵ ୢ୶

ൌ െ ............................................. (5.4)

ୢ୲ ଢ଼౮Ȁ౩ ୢ୲

ଵ ୢ୶

െ ൌ െ

ଢ଼౮Ȁ౩ ୢ୲

!

ଵ

ൌ െ ሺ୲ െ ሻ

ଢ଼౮Ȁ౩

$ !

The growth of microbial cells include four major phases namely lag phase,

exponential phase, stationary phase and the decline phase, which had been previously

described in chapter II. An emphasis on the exponential or logarithmic phase had been

given in this particular work, since the production of the product, exopolysaccharide, is

found maximum and directly proportional to the microbial cell growth.

Various models had been used for studying growth kinetics. The models are being

mentioned under this topic.

The simplest cell growth model is the Malthusian model commonly known as

exponential law. In a batch fermentation operated under ideal conditions the

concentration and weight of biomass increases proportionally with increase in time.

Hence it could be said that all the cells have the same probability to multiply. Thus the

overall rate of biomass formation is proportional to the biomass itself. For a batch system,

this is equivalent to

ୢ୶

ൌ Ɋ .................................... (5.7)

ୢ୲

where x is the mass of cells per unit volume, µ is the proportionality constant known as

'specific growth rate' (h-1) and t is the time (h).

Here as the time increases, cell mass will also increase. On integrating (1) gives

The demerit of this model is that it ignores the substrate needed for its growth and

that the resources are constrained.

Contois model is another unstructured model that depends upon substrate and cell

concentration which is given as

ୱ

Ɋ ൌ Ɋ୫ୟ୶

୶ାୱ

& !

Moser model relies upon the growth rate and substrate concnetration and the two

parameter model is depicted as

ିଵ

Ɋ ൌ Ɋ୫ୟ୶ ൫ͳ ୱ ି ൯ ' !

Tessier model is another growth kinetic model that also rely upon the substrate

concentration and is explained by,

ୱ

Ɋ ൌ Ɋ୫ୟ୶ ൫ͳ െ ି ൗୱ ൯ .......................... (5.11)

Moser and Tessier model render algebraic solution of the growth equations much

more difficult than the Monod model.

%

5.2.5 Verlhurst model

Verlhurst kinetic model depends on the cell concentration. This model has two

kinetic constants of maximum specific growth rate (µmax) and maximum cell

concentration (xmax). Verlhurst model is expressed by the following equation

ஜౣ౮

Ɋ ൌ Ɋ୫ୟ୶ െ

୶ౣ౮

" !

Andrew model is proposed by the following equation for the growth dependent

which incorporate substrate inhibition.

ஜౣ౮

Ɋ ൌ ౩ൗ ୱ

............... (5.13)

ቂቀଵି ୱቁቀଵା ൗ౩ ቁቃ

The commonly used unstructured kinetic models to evaluate cell growth are

Monod and Logistic models.

The concepts in microbial growth kinetics have been mainly analysed by semi-

empirical model proposed by Monod (1949). The Monod model introduces the concept

of a growth controlling substrate, called limiting substrate which indicates that the

microbial growth rate rely upon the actual concentration of a particular metabolite. There

exists a relationship between the exhaustion of the limiting substrate and the end of

growth, hence the Monod model may be considered deterministic. The relationship is

described as,

ஜౣ౮ ୱ

Ɋ ൌ ............................................. (5.14)

౩ ାୱ

where µmax is the maximum growth rate achievable and Ks is the limiting substrate

concentration when the specific growth rate is equal to the half the maximum specific

growth rate when µ = µmax/2.

(

5.2.8 Logistic model

The model which could be appropriate for the present research is the logistic

model. Verlhurst in 1844 and Pearl and Reed in 1920 contributed to a theory which

included an inhibiting factor to population growth. Assuming that inhibition is

proportional to x2, they used

ୢ୶ ୶

ൌ ሺͳ െ ሻ

ୢ୲ ୶౩

* !

where x0 is the initial biomass concentration (gL-1) and t is the time (h). The logistic

curve is sigmoidal and leads to a stationary population of size, xs +

, -

the given time. When the cell mass reaches the stationary phase, the growth rate ceases.

A gradual decrease is observed in the late exponential phase or when the cells near the

stationary phase. Advantage of this model is that it facilitates the exponential phase and

endogenous metabolic phase. It also has its own drawback that it fails to predict a decline

phase after the stationary phase.

The type of kinetic description that may be employed for product formation by

cell population parallel those used to describe cell population growth. The simplest types

of product formation kinetics arise where there is a simple stoichiometric connection

between product formation and substrate utilization or cell growth. The rate of production

of the product parallels the rate of cell growth. Substrate concentration falls continuously

as fermentation proceeds. Such product formation kinetics is sometimes called as growth

associated. In many fermentation, involving secondary metabolite, significant product

formation does not occur until relatively late in a batch cultivation, that is, production

happens in the late exponential phase or during the stationary phase. Production rate is

proportional to cell concentration rather than growth rate. This kind of kinetics is referred

to as non growth associated model.

)

A typical and widely used product kinetic model is Luedeking- Piret model

(1959), which is an unstructured approach contributed to both growth and non growth

associated phenomena for product formation. This model was originally developed for

the formation of lactic acid by Lactobacillus delbruckeii and the equation is given as

rfp = fx . . . . . . . . . / / / / / / / / / / / / / 0 1 / 2 1 3

where rfp is the rate of product formation, rfx is the rate of biomass formation and 4 and 5

are the kinetic constants of Luedeking- Piret model. This two-parameter expression has

proven extremely useful and versatile in fitting product formation data from different

fermentations. This is an expected kinetic form when the product is the result of energy

yielding metabolism.

According to this model, the product formation rate depends linearly upon the

growth rate and the cell concentration

ୢ ୢ୶

ൌ Ƚ Ⱦ ...................................... (5.16)

ୢ୲ ୢ୲

The Luedeking- 6 7 8 9 : ; 7 < 9 : 7 = > ? 8 ? @ 9 : 9 8 A B C ? < D E D 9 > 9 < D F > G < ? < D H ? 8 I J 7 : K : K 9

L L

ౚౌ

ቀ ቁୱ୲ୟ୲୧୭୬ୟ୰୷୮୦ୟୱୣ

ౚ౪

Ⱦൌ ................................ (5.17)

୶౩

ܻȀ௫ ,

L L

ୟୱୱ୭୮୰୭ୢ୳ୡ୲୭୰୫ୣୢ ିబ

Ȁ୶ ൌ ൌ ൌ ................... (5.18)

ୟୱୱ୭ୡୣ୪୪୭୰୫ୣୢ ୶ ୶ି୶బ

୶ ୶

ሺሻ െ ሺͲሻ െ Ⱦ ቀ ౩ ቁቂͳ െ బ ൫ͳ െ ୩୲ ൯ቃ ൌ Ƚሺ୲ െ ሻ .............. (5.19)

୩ ୶ ౩

L

vs. (xt x0).

S

Cell growth, substrate utilisation and product formation were examined and

simulated with the experimental data of EPS yield by B.subtilis and P.fluorescens, using

cane molasses and rice bran respectively. The logistic equation was used for the cellular

growth kinetic study and Luedeking Piret model for substrate consumption and product

formation studies. The simulation of the experiment was carried out using MATLAB

(v.7.10.0.0499, The Mathworks, USA) software. The MATLAB program is listed in

Appendix I and II. The kinetic parameter 'k' of logistic model was obtained using curve

fitting (cftool) tool kit of the same software and the high R2 values represented that the

equation fit the experiment (Sivaprakash et. al., 2011a). Using the obtained 'k' values for

each biological system, the kinetic constants of Luedeking- Piret model, α and β, were

evaluated (Table 5.1). The model was analysed using MS- Excel by graphical method.

The yield coefficient Yx/s, for substrate utilisation rate, was also obtained by graphical

method. Predicted values of this model, which were consistent with the observed values,

were obtained by solving the differential equations by Runge K F : : ? T A < F @ 9 8 7 = ? O

integration using ODE23 solver, in same software, which is given in Appendix III

(Sivaprakash et. al., 2011b). The theoretical and predicted values obtained are tabulated

in Table 5.2 and Table 5.3.

Substrate

Product formation

Organism Consumption

k Error Error

R2 Yx/s

(h-1)

C E

% %

B.subtilis 0.09126 0.9829 10.22 6.63 0.5 0.0808 3.05

P.fluorescens 0.08422 0.9852 5.5 5.45 0.6 0.114 7.47

Q R R

The high R2 and prediction values denote that the model is highly suitable for

exopolysaccharide production with a minimal error of 4.52 % and 9.91% from Bacillus

subtilis and Pseudomonas fluorescens. Figures 5.3 5.4, 5.5, 5.6, 5.7 and 5.8, illustrate the

experimental and predicted values for each variable S

product formed at a given time.

Table 5.2 Experimental and predicted values of cell mass concentration, substrate

utilization and product formation of B.subtilis

Time, t Concentration, Consumption Formation, P

(h) x (gL-1) (gL-1) (gL-1)

Experimental Predicted Experimental Predicted Experimental Predicted

0 1.893 1.893 1.467 1.467 0 0

6 3.432 2.944195 1.318 1.364143 0.61 0.579208

12 3.989 4.337211 1.304 1.22784 1.281 1.34676

18 6.843 5.971837 0.939 1.067897 2.37 2.247439

24 8.761 7.636388 0.876 0.905025 2.972 3.164607

30 8.834 9.104527 0.872 0.761371 3.825 3.973551

36 9.743 10.24598 0.725 0.649683 4.582 4.60249

42 9.955 11.04985 0.73 0.571027 4.672 5.045422

48 11.401 11.58243 0.526 0.518915 4.973 5.338877

54 11.593 11.90793 0.521 0.487065 5.389 5.518227

60 11.932 12.1016 0.473 0.468115 5.421 5.624939

66 11.992 12.21622 0.461 0.4569 5.561 5.688096

72 12.364 12.27992 0.428 0.450668 5.731 5.723191

78 12.364 12.31684 0.419 0.447055 5.731 5.743534

84 12.364 12.33759 0.418 0.445024 5.731 5.754972

90 12.364 12.34906 0.417 0.443902 5.731 5.761287

96 12.364 12.35559 0.415 0.443263 5.731 5.764889

Q R U

14

12

10

4

Experimental

2 Simulated

0

0 20 40 60 80 100 120

Time (h)

Figure 5.3 Comparative chart showing experimental and predicted values of cell

concentration of B.subtilis

1.6

Experimental

1.4

Simulated

Substrate Consumption (g/L)

1.2

0.8

0.6

0.4

0.2

0

0 20 40 60 80 100 120

Time (h)

substrate consumption of B.subtilis

Q R V

7

Product (g/L)

4

2 Experimental

Simulated

1

0

0 20 40 60 80 100 120

Time (h)

product concentration of B.subtilis

Table 5.2 Experimental and predicted values of cell mass concentration, substrate

utilization and product formation of Pseudomonas fluorescens

Time, Concentration, Consumption Formation, P

t -1

x (gL ) -1

(gL ) (gL-1)

(h) Experimental Predicted Experimental Predicted Experimental Predicted

0 1.036 1.036 1.897 1.897 0 0

6 1.428 1.593345 1.768 1.795665 0.352 0.334407

12 2.014 2.358844 1.642 1.656483 0.702 0.793706

18 2.414 3.322078 1.521 1.481349 1.275 1.371647

24 3.986 4.408413 1.502 1.283834 1.917 2.023448

30 5.002 5.492006 1.207 1.086817 2.474 2.673603

36 6.789 6.448431 0.924 0.912922 3.381 3.247458

42 7.136 7.206297 0.816 0.775128 3.704 3.702178

48 8.771 7.756798 0.608 0.675037 4.861 4.032479

54 8.771 8.131481 0.606 0.606912 4.861 4.257289

60 8.771 8.375118 0.605 0.562615 4.861 4.403471

66 8.771 8.528178 0.604 0.534786 4.861 4.495307

Q R W

10

9

8

7

Cell mass (g/L) 6

5

4

3 Experimental

2 Simulated

1

0

0 10 20 30 40 50 60 70

Time (h)

Figure 5.6 Comparative chart showing experimental and predicted values of cell

concentration of P.fluorescens

2

1.8

Experimental

Substrate Consumption (g/L)

1.6 Simulated

1.4

1.2

1

0.8

0.6

0.4

0.2

0

0 10 20 30 40 50 60 70

Time (h)

Figure 5.7 Comparative chart showing experimental and predicted values of cell

substrate consumption of P.fluorescens

Q U X

6

Product (g/L)

4

0

0 10 20 30 40 50 60 70

Time (h)

product concentration of P.fluorescens

The utilization rate was well predicted and found to be the most suited model

for the exopolysaccharide production from cane molasses and rice bran respectively with

a minimal error of 6.63% and 5.45% for Bacillus subtilis and Pseudomonas fluorescens

respectively. Similarly, the predicted and theoretical values were found to be in

agreement with the experimental data of the exopolysaccharide formation kinetics with a

minimum error of 3.05% and 7.47% for Bacillus subtilis and Pseudomonas fluorescens

respectively.

The overall trend of the simulated or the predicted values, obtained using the

implemented models in accordance with the experimental data for B.subtilis and

P.fluorescens is depicted in Figure 5.9. The studies confirmed that the Logistic and

Luedeking Piret models befitted accurately and explained adequately the mechanisms of

cell growth, substrate consumption and product formation in B.subtilis and P.

fluorescens.

Q U Q

14 2

1.8

12

Product (g/L) 1.6

10

1.4

1.2

8

1

Cell mass (g/L)

6

0.8

4 0.6

0.4

2

0.2

0 0

0 20 40 60 80 100 120

Time (h)

Figure 5.9 Overall experimental and predicted values of Logistic and Luedeking

Piret models for Bacillus subtilis and Pseudomonas fluorescens (cell mass of B.

subtilis, experimental and predicted ; product formation by B. subtilis,

experimental and predicted ; substrate consumed by B. subtilis,

experimental and predicted ; cell mass of P. fluorescens, experimental

and predicted ; product formation by P. fluorescens experimental and

predicted ; substrate consumed by P. fluorescens experimental and

predicted)

The viscosity measurements were performed for the cultures after 24 h till 120h to

check their viscosity. The viscosities and relative viscositites were found to be constant

after certain period (Fig.5.10). In case of B.subtilis, the relative viscosity was constant

after 72h, whereas for culture broth of P.fluorescens, the relative viscosity stayed the

same after 48h. The trends of both cultures exhibited a Newtonian behaviour.

Q U Y

1.02

1

0.98

Relative viscosity

0.96

0.94

0.92

0.9

B.subtilis

0.88

P.fluorescens

0.86

0.84

0 20 40 60 80 100 120

Time (h)

Figure 5.10 Relative viscosity of the two culture filtrate with EPS

shear stress. Dilute polymer solutions exhibit approximately Newtonian behavior at low

shear rates. For Newtonian fluids, shear stress is directly proportional to the shear rate,

? < D : K 9 > 8 G > G 8 : 7 G < ? O 7 : I = G < A : ? < : 7 A H 7 A = G A 7 : I B [ / The study is in agreement with Al Nahas

et al (2011) who reported the similar trend for EPS produced by Pseudoalteromonas sp.

As the cells grew, the viscosity kept increasing. The polymeric solution exhibited a non-

Newtonian behavior. This showed that the maximum EPS was found to be produced

during the exponential phase and after the stationary phase the production remained

stable.

Q U Z

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