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IOSR Journal of Mathematics (IOSR-JM)
e-ISSN: 2278-5728,p-ISSN: 2319-765X, Volume 7, Issue 2 (Jul. - Aug. 2013), PP 19-22
www.iosrjournals.org
www.iosrjournals.org 19 | Page
Parametric sensitivity analysis of a mathematical model of
facultative mutualism
Enu Ekaka-a, 1
E.C. Nwachukwu, N.M. Nafo, I.A. Agwu,
Department of Mathematics and Computer Science, Rivers State University of Science and Technology, Port
Harcourt, Rivers State, Nigeria
Department of Mathematics and Statistics, University of Port Harcourt, Nigeria
Department of Mathematics and Computer Science, Rivers State University of Science and Technology, Port
Harcourt, Nigeria
Department of Mathematics, Abia State Polytechnic, Aba, Abia State, Nigeria
Abstract: The complex dynamics of facultative mutualism is best described by a system of continuous non-linear
first order ordinary differential equations. The methods of 1-norm, 2-norm, and infinity-norm will be used to
quantify and differentiate the different forms of the sensitivity of model parameters. These contributions will be
presented and discussed.
I. Introduction
According to Morin (2002), competition and predation have been observed as two dominant types of
interaction between species. In the literatures of mathematical ecology or mathematical biology, competition is the
most recognised type of interaction while mutualism despite its relevance seems to be not frequently mentioned in
most research agenda. Having conducted a detailed literature review on the topic of the dynamics of transitions
between population interactions, using a non-linear α-function, we found that the application of parametric
sensitivity analysis of the model parameters for this important model is rarely introduced as a method of studying
the importance of the model parameters of such model. We are motivated to implement the tool of parametric
sensitivity analysis because it is the first stage in the process of developing a deterministic model which describes
the dynamics of two interacting populations. To the best of our knowledge, we are yet to see any reported
mathematical analysis which involves the utilization of a numerical parametric sensitivity analysis. This method is
expected to estimate and classify the sensitivity of model parameters which were proposed by Hernandez (1998). In
the next section, we will present the mathematical formulation of the model on which our analysis id based.
II. Mathematical Formulation
Following Hernandez (1998), we consider the following complex system of two continuous nonlinear first
order ordinary differential equations of the following mathematical structure:
𝑑𝑁1
𝑑𝑡
= 𝑟1 𝑁1 1 −
𝑁1
𝐾1
+
𝑏1 𝑁2 − 𝑐1 𝑁2
2
1 + 𝑑1 𝑁2
2
𝑁2
𝐾1
𝑑𝑁2
𝑑𝑡
= 𝑟2 𝑁2 1 −
𝑁2
𝐾2
+
𝑏2 𝑁1 − 𝑐2 𝑁1
2
1 + 𝑑2 𝑁1
2
𝑁1
𝐾2
Here, 𝑁1 and 𝑁2 are populations of specie 1 and specie 2 respectively, 𝑟1 and 𝑟2are growth rates of specie 1 and
specie 2 respectively, 𝐾1 and 𝐾2are carrying capacities of specie 1 and specie 2 respectively.The parameters
𝑏1, 𝑏2, 𝑐1, 𝑐2, 𝑑1and𝑑2modify the general shape and represent the environmentalinfluence on αijthat is, the
“exogenous effect”.
This facultative model (Hernandez (1998)) is driven by thirteen (13) parameters. To conduct a realistic and
systematic parametric sensitivity analysis of this model, we have proposed to study the sensitivity analysis of the
two intrinsic growth rates (r1 and r2), carrying capacities (K1 and K2) and the initial conditions (N1(0) and N2(0)) for
obvious ecological or biological reasons. In the next section, we will describe the key relevant materials and
methods which will enable us to solve this problem numerically.
Submitted Date 30 May 2013 Accepted Date: 06 June 2013
Parametric sensitivity analysis of a mathematical model of facultative mutualism
www.iosrjournals.org 20 | Page
III. Materials and Methods
Following Ford et al (2010) and Hernandez (1998), these researchers in mathematical biology have
identified similar types of interactions between species such as mutualism, commensalism, parasitism or predation
and competition. The method of Ford et al (2010) was based on using the biological ideas to construct a model for
two competing plant species in a harsh climate whereas the model of Hernandez (1998) simply describes in great
detail the undergoing transitions between different types of partially positive interaction hereby called facultative
and obligate mutualism which is consistent with the classification of Morin (2002, 187 – 189).
In our recent literature review, the current state of the spread of knowledge clearly shows that the work of
Hernandez (1998) has been popularly cited by 32 other researchers. However, none of these researchers has
attempted to study the parametric sensitivity analysis of the facultative model.Hernandez (1998) considered an
interaction where both α-functions are of the α(+, -) type and reported that at low densities, the association is
mutualistic but at higher densities the same association was considered to be competitive. It was also reported in the
same paper that depending on the threshold value where α12 and α21 are equal to zero, immediate density values can
be experimentally determined for a situation where a parasitic interaction can also occur. Part of developing this
model involves assuming that both species can exist either alone or in association. The approximate model
formulation was constructed on the basis of these simplifying assumptions. The method of sensitivity analysis which
we have used in this study has been adapted from recent research reports of Ekaka-a (2009), Ekaka-a et al (2012)
and Nwachukwu and Ekaka-a (2013). A brief sketch of this method is as follows:
STEP I: Code the given system of continuous non-linear first order ordinary differential equation in a Matlab
programing language.
STEP II: Modify and code a similar program which is used for a variation of a single parameter one-at-a-time while
other model parameters are fixed.
STEP III: Design an appropriate ODE45 Runge-Kutta scheme which will simulate the program in step I and step II
STEP IV: Use the program in step III to calculate the 1-norm, 2-norm, and infinity norm of three solution
trajectories in the same manner, use the same program to calculate the three popular norms of the differences of the
solution trajectories.
STEP V: Based on the original parameter in the step I, calculate the cumulative percentage effect on the solution
trajectories due to variation of each chosen parameter at a time when other parameters are fixed.
STEP VI: Interprete the result quantitatively. That is, the parameter which when varied a little and produces the
biggest cumulative effect on the solution trajectories is called a most sensitive.
IV. Results andDiscussion
In this section, we will present and discuss our results which we have achieved in this study. In this section,
the notation CV stands for the popular statistical coefficient of variation.
Table 1 Percentage variations of r1
variations
mn
0.015 0.03 0.045 0.06 0.075
1-norm 10.5213 5.3085 3.5391 2.6518 2.1221
2-norm 10.9065 7.5868 6.0936 5.1905 4.5654
∞-norm 32.5020 30.7169 29.1662 27.6902 26.5351
Table 2 Percentage variations of r2
variations
mn
0.015 0.030 0.045 0.06 0.075
1-norm 18.5029 9.1881 6.0497 4.4794 3.5403
2-norm 22.8126 15.5895 12.3174 10.3293 8.9562
∞-norm 98.7772 88.7117 81.2441 74.9255 69.5011
Table 3 Percentage variations of K1
variations
mn
0.01 0.02 0.03 0.04 0.05
1-norm 65.2987 64.9995 64.6973 64.3887 64.0780
2-norm 47.5824 47.3460 47.1080 46.8654 46.6221
∞-norm 54.2792 54.0124 53.7196 53.4540 53.1737
Parametric sensitivity analysis of a mathematical model of facultative mutualism
www.iosrjournals.org 21 | Page
Table 4 Percentage variations of K2
variations
mn
0.01 0.02 0.03 0.04 0.05
1-norm 65.4713 65.1745 64.8735 64.5666 64.2571
2-norm 47.5803 47.3472 47.1114 46.8715 46.6304
∞-norm 62.0988 61.9169 61.6954 61.4953 61.2576
Table 5 Percentage variations of IC1
variations
mn
0.04 0.08 0.12 0.16 0.20
1-norm 0.5121 0.5079 0.5028 0.5022 0.5013
2-norm 5.5154 5.4601 5.4051 5.3500 5.2949
∞-norm 74.1298 73.5982 73.0003 72.3749 71.7193
Table 6 Percentage variations of IC2
variations
mn
0.1 0.2 0.3 0.4 0.5
1-norm 1.3768 1.3709 1.3681 1.3499 1.3443
2-norm 13.8051 13.6737 13.5407 13.4063 13.2710
∞-norm 117.6685 116.9752 116.1566 115.2819 114.3127
Table 7 results of coefficient of variation for model parameters
parameter r1 CV parameter K1 CV parameter IC1 CV
1-norm 0.9640 1-norm 0.0075 1-norm 0.0091
2-norm 0.4310 2-norm 0.0081 2-norm 0.0162
∞-norm 0.0838 ∞-norm 0.0082 ∞-norm 0.0132
parameter r2 CV parameter K2 CV parameter IC2 CV
1-norm 1.0038 1-norm 0.0074 1-norm 0.0104
2-norm 0.4678 2-norm 0.0080 2-norm 0.0157
∞-norm 0.1481 ∞-norm 0.0054 ∞-norm 0.0115
V. Discussion of results
From our results which have been presented in the previous section, we observe that the two carrying
capacities can be classified as the most sensitive parameters using the 1-norm and 2-norm estimated sensitivity
values while the two initial conditions can be classified as relatively least sensitive parameters. However, the ∞-
norm estimated value shows that the second initial condition is clearly a more sensitive parameter than any of the
other five model parameters. In the context of measuring the best estimate of sensitivity, we can clearly see that the
infinity norm is the best estimate for measuring the sensitivity of model parameters such as r1, r2, and K2 while the 1-
norm is the best estimate for measuring the sensitivity of model parameters such as K1 and the initial conditions.
VI. Conclusion
In terms of further parameter estimation to minimize uncertainty in model prediction, carrying capacities
and the growth rates should be targeted for the purpose of model re-validation. In this study, the initial conditions in
the main should be considered as rough estimates. In our further research, we propose to study the sensitivity of
other seven (7) parameters which we have not considered in this present study.
References
[1]. Ekaka-a E. N. (2009): Computational and Mathematical Modelling of Plant Species interactions in a Harsh Climate. PhD Thesis, Dept. of
Mathematics, The university of Liverpool and The University of Chester, United Kingdom.
[2]. Ford Neville J, Lumb Patricia M, Ekaka-a Enu (2010): Mathematical modelling of plant species interactions in a harsh climate, Journal of
Computational and Applied Mathematics, Vol. 234, pp. 2732-2744.
[3]. Hernandez MJ (1998): Dynamics of transitions between population interactions: a nonlinear interaction alpha-function defined,
Proceedings of the Royal Society B Biological Sciences London 1998 265, 1433-1440..
Parametric sensitivity analysis of a mathematical model of facultative mutualism
www.iosrjournals.org 22 | Page
[4]. Nwachukwu E. C. and E. N. Ekaka-a (2013): Sensitivity Analysis using a partially coupled system of differential equations without delay,
Journal of Mathematics and System Science (ISSN 2159-5291, USA).
[5]. Morin P.J (2002), Community Ecology, Blackwell Publishing Company, United Kingdom.

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Parametric sensitivity analysis of a mathematical model of facultative mutualism

  • 1. IOSR Journal of Mathematics (IOSR-JM) e-ISSN: 2278-5728,p-ISSN: 2319-765X, Volume 7, Issue 2 (Jul. - Aug. 2013), PP 19-22 www.iosrjournals.org www.iosrjournals.org 19 | Page Parametric sensitivity analysis of a mathematical model of facultative mutualism Enu Ekaka-a, 1 E.C. Nwachukwu, N.M. Nafo, I.A. Agwu, Department of Mathematics and Computer Science, Rivers State University of Science and Technology, Port Harcourt, Rivers State, Nigeria Department of Mathematics and Statistics, University of Port Harcourt, Nigeria Department of Mathematics and Computer Science, Rivers State University of Science and Technology, Port Harcourt, Nigeria Department of Mathematics, Abia State Polytechnic, Aba, Abia State, Nigeria Abstract: The complex dynamics of facultative mutualism is best described by a system of continuous non-linear first order ordinary differential equations. The methods of 1-norm, 2-norm, and infinity-norm will be used to quantify and differentiate the different forms of the sensitivity of model parameters. These contributions will be presented and discussed. I. Introduction According to Morin (2002), competition and predation have been observed as two dominant types of interaction between species. In the literatures of mathematical ecology or mathematical biology, competition is the most recognised type of interaction while mutualism despite its relevance seems to be not frequently mentioned in most research agenda. Having conducted a detailed literature review on the topic of the dynamics of transitions between population interactions, using a non-linear α-function, we found that the application of parametric sensitivity analysis of the model parameters for this important model is rarely introduced as a method of studying the importance of the model parameters of such model. We are motivated to implement the tool of parametric sensitivity analysis because it is the first stage in the process of developing a deterministic model which describes the dynamics of two interacting populations. To the best of our knowledge, we are yet to see any reported mathematical analysis which involves the utilization of a numerical parametric sensitivity analysis. This method is expected to estimate and classify the sensitivity of model parameters which were proposed by Hernandez (1998). In the next section, we will present the mathematical formulation of the model on which our analysis id based. II. Mathematical Formulation Following Hernandez (1998), we consider the following complex system of two continuous nonlinear first order ordinary differential equations of the following mathematical structure: 𝑑𝑁1 𝑑𝑡 = 𝑟1 𝑁1 1 − 𝑁1 𝐾1 + 𝑏1 𝑁2 − 𝑐1 𝑁2 2 1 + 𝑑1 𝑁2 2 𝑁2 𝐾1 𝑑𝑁2 𝑑𝑡 = 𝑟2 𝑁2 1 − 𝑁2 𝐾2 + 𝑏2 𝑁1 − 𝑐2 𝑁1 2 1 + 𝑑2 𝑁1 2 𝑁1 𝐾2 Here, 𝑁1 and 𝑁2 are populations of specie 1 and specie 2 respectively, 𝑟1 and 𝑟2are growth rates of specie 1 and specie 2 respectively, 𝐾1 and 𝐾2are carrying capacities of specie 1 and specie 2 respectively.The parameters 𝑏1, 𝑏2, 𝑐1, 𝑐2, 𝑑1and𝑑2modify the general shape and represent the environmentalinfluence on αijthat is, the “exogenous effect”. This facultative model (Hernandez (1998)) is driven by thirteen (13) parameters. To conduct a realistic and systematic parametric sensitivity analysis of this model, we have proposed to study the sensitivity analysis of the two intrinsic growth rates (r1 and r2), carrying capacities (K1 and K2) and the initial conditions (N1(0) and N2(0)) for obvious ecological or biological reasons. In the next section, we will describe the key relevant materials and methods which will enable us to solve this problem numerically. Submitted Date 30 May 2013 Accepted Date: 06 June 2013
  • 2. Parametric sensitivity analysis of a mathematical model of facultative mutualism www.iosrjournals.org 20 | Page III. Materials and Methods Following Ford et al (2010) and Hernandez (1998), these researchers in mathematical biology have identified similar types of interactions between species such as mutualism, commensalism, parasitism or predation and competition. The method of Ford et al (2010) was based on using the biological ideas to construct a model for two competing plant species in a harsh climate whereas the model of Hernandez (1998) simply describes in great detail the undergoing transitions between different types of partially positive interaction hereby called facultative and obligate mutualism which is consistent with the classification of Morin (2002, 187 – 189). In our recent literature review, the current state of the spread of knowledge clearly shows that the work of Hernandez (1998) has been popularly cited by 32 other researchers. However, none of these researchers has attempted to study the parametric sensitivity analysis of the facultative model.Hernandez (1998) considered an interaction where both α-functions are of the α(+, -) type and reported that at low densities, the association is mutualistic but at higher densities the same association was considered to be competitive. It was also reported in the same paper that depending on the threshold value where α12 and α21 are equal to zero, immediate density values can be experimentally determined for a situation where a parasitic interaction can also occur. Part of developing this model involves assuming that both species can exist either alone or in association. The approximate model formulation was constructed on the basis of these simplifying assumptions. The method of sensitivity analysis which we have used in this study has been adapted from recent research reports of Ekaka-a (2009), Ekaka-a et al (2012) and Nwachukwu and Ekaka-a (2013). A brief sketch of this method is as follows: STEP I: Code the given system of continuous non-linear first order ordinary differential equation in a Matlab programing language. STEP II: Modify and code a similar program which is used for a variation of a single parameter one-at-a-time while other model parameters are fixed. STEP III: Design an appropriate ODE45 Runge-Kutta scheme which will simulate the program in step I and step II STEP IV: Use the program in step III to calculate the 1-norm, 2-norm, and infinity norm of three solution trajectories in the same manner, use the same program to calculate the three popular norms of the differences of the solution trajectories. STEP V: Based on the original parameter in the step I, calculate the cumulative percentage effect on the solution trajectories due to variation of each chosen parameter at a time when other parameters are fixed. STEP VI: Interprete the result quantitatively. That is, the parameter which when varied a little and produces the biggest cumulative effect on the solution trajectories is called a most sensitive. IV. Results andDiscussion In this section, we will present and discuss our results which we have achieved in this study. In this section, the notation CV stands for the popular statistical coefficient of variation. Table 1 Percentage variations of r1 variations mn 0.015 0.03 0.045 0.06 0.075 1-norm 10.5213 5.3085 3.5391 2.6518 2.1221 2-norm 10.9065 7.5868 6.0936 5.1905 4.5654 ∞-norm 32.5020 30.7169 29.1662 27.6902 26.5351 Table 2 Percentage variations of r2 variations mn 0.015 0.030 0.045 0.06 0.075 1-norm 18.5029 9.1881 6.0497 4.4794 3.5403 2-norm 22.8126 15.5895 12.3174 10.3293 8.9562 ∞-norm 98.7772 88.7117 81.2441 74.9255 69.5011 Table 3 Percentage variations of K1 variations mn 0.01 0.02 0.03 0.04 0.05 1-norm 65.2987 64.9995 64.6973 64.3887 64.0780 2-norm 47.5824 47.3460 47.1080 46.8654 46.6221 ∞-norm 54.2792 54.0124 53.7196 53.4540 53.1737
  • 3. Parametric sensitivity analysis of a mathematical model of facultative mutualism www.iosrjournals.org 21 | Page Table 4 Percentage variations of K2 variations mn 0.01 0.02 0.03 0.04 0.05 1-norm 65.4713 65.1745 64.8735 64.5666 64.2571 2-norm 47.5803 47.3472 47.1114 46.8715 46.6304 ∞-norm 62.0988 61.9169 61.6954 61.4953 61.2576 Table 5 Percentage variations of IC1 variations mn 0.04 0.08 0.12 0.16 0.20 1-norm 0.5121 0.5079 0.5028 0.5022 0.5013 2-norm 5.5154 5.4601 5.4051 5.3500 5.2949 ∞-norm 74.1298 73.5982 73.0003 72.3749 71.7193 Table 6 Percentage variations of IC2 variations mn 0.1 0.2 0.3 0.4 0.5 1-norm 1.3768 1.3709 1.3681 1.3499 1.3443 2-norm 13.8051 13.6737 13.5407 13.4063 13.2710 ∞-norm 117.6685 116.9752 116.1566 115.2819 114.3127 Table 7 results of coefficient of variation for model parameters parameter r1 CV parameter K1 CV parameter IC1 CV 1-norm 0.9640 1-norm 0.0075 1-norm 0.0091 2-norm 0.4310 2-norm 0.0081 2-norm 0.0162 ∞-norm 0.0838 ∞-norm 0.0082 ∞-norm 0.0132 parameter r2 CV parameter K2 CV parameter IC2 CV 1-norm 1.0038 1-norm 0.0074 1-norm 0.0104 2-norm 0.4678 2-norm 0.0080 2-norm 0.0157 ∞-norm 0.1481 ∞-norm 0.0054 ∞-norm 0.0115 V. Discussion of results From our results which have been presented in the previous section, we observe that the two carrying capacities can be classified as the most sensitive parameters using the 1-norm and 2-norm estimated sensitivity values while the two initial conditions can be classified as relatively least sensitive parameters. However, the ∞- norm estimated value shows that the second initial condition is clearly a more sensitive parameter than any of the other five model parameters. In the context of measuring the best estimate of sensitivity, we can clearly see that the infinity norm is the best estimate for measuring the sensitivity of model parameters such as r1, r2, and K2 while the 1- norm is the best estimate for measuring the sensitivity of model parameters such as K1 and the initial conditions. VI. Conclusion In terms of further parameter estimation to minimize uncertainty in model prediction, carrying capacities and the growth rates should be targeted for the purpose of model re-validation. In this study, the initial conditions in the main should be considered as rough estimates. In our further research, we propose to study the sensitivity of other seven (7) parameters which we have not considered in this present study. References [1]. Ekaka-a E. N. (2009): Computational and Mathematical Modelling of Plant Species interactions in a Harsh Climate. PhD Thesis, Dept. of Mathematics, The university of Liverpool and The University of Chester, United Kingdom. [2]. Ford Neville J, Lumb Patricia M, Ekaka-a Enu (2010): Mathematical modelling of plant species interactions in a harsh climate, Journal of Computational and Applied Mathematics, Vol. 234, pp. 2732-2744. [3]. Hernandez MJ (1998): Dynamics of transitions between population interactions: a nonlinear interaction alpha-function defined, Proceedings of the Royal Society B Biological Sciences London 1998 265, 1433-1440..
  • 4. Parametric sensitivity analysis of a mathematical model of facultative mutualism www.iosrjournals.org 22 | Page [4]. Nwachukwu E. C. and E. N. Ekaka-a (2013): Sensitivity Analysis using a partially coupled system of differential equations without delay, Journal of Mathematics and System Science (ISSN 2159-5291, USA). [5]. Morin P.J (2002), Community Ecology, Blackwell Publishing Company, United Kingdom.