Mathematical Modeling

Prof.Lakshmi Burra
 Models for
Population Growth

In this section, we will:

Investigate differential equations
used to model population growth.
NATURAL GROWTH

One of the models for population growth is

based on the assumption that the population
grows at a rate proportional to the size of the
population:
NATURAL GROWTH

In general, it seems reasonable that

the growth rate should be proportional
to the size.
LAW OF NATURAL GROWTH Equation 1

In general, if P(t) is the value of a quantity y

at time t and, if the rate of change of P with
respect to t is proportional to its size P(t) at
any time, then

where k is a constant.

 This is sometimes called the law of natural growth.

LAW OF NATURAL GROWTH

If k is positive,
the population increases.

If k is negative, it decreases.
 LAW OF NATURAL GROWTH

Equation 1 is a separable differential equation.Hence, we can solve

it by usual methods

where A (= ±eC or 0) is an arbitrary constant.

LAW OF NATURAL GROWTH

To see the significance of the constant A,

we observe that:

P(0) = Aek·0 = A

Thus, A is the initial value of the function.

LAW OF NATURAL GROWTH Equation 2

The solution of the initial-value problem

is:
LAW OF NATURAL GROWTH

Another way of writing Equation 1 is:

 This says that the relative growth rate (the growth rate
divided by the population size) is constant.

 Then, Equation 2 says that a population with

constant relative growth rate must grow exponentially.
LOGISTIC MODEL

The simplest expression for the

relative growth rate that incorporates
these assumptions is:
LOGISTIC DIFFERENTIAL EQN. Equation 4

Multiplying by P, we obtain the model

for population growth known as the logistic
differential equation:
LOGISTIC DIFFERENTIAL EQN.

Notice from Equation 4 that:

 If P is small compared with K, then P/K is close to 0,

and so dP/dt ≈ kP.

 If P → K (the population approaches its carrying

capacity), then P/K → 1, so dP/dt → 0.
LOGISTIC DIFFERENTIAL EQN.

From Equation 4, we can deduce

information about whether solutions
increase or decrease directly.
LOGISTIC DIFFERENTIAL EQN.

If the population P lies between 0 and K,

the right side of the equation is positive.

 So, dP/dt > 0 and the population increases.

If the population exceeds the carrying

capacity (P > K), 1 – P/K is negative.

 So, dP/dt < 0 and the population decreases.

LOGISTIC DIFFERENTIAL EQN.

Let’s start our more detailed analysis

of the logistic differential equation by
looking at a direction field.
LOGISTIC DIFFERENTIAL EQN. Example 1

Notice that solution curves that start:

 Below P = 1000 are increasing.

 Above P = 1000 are decreasing.
LOGISTIC DIFFERENTIAL EQN.

The logistic equation 4 is separable.

So, we can solve it explicitly

MODELS FOR POPULATION GROWTH

The Law of Natural Growth and the logistic

differential equation are not the only equations
that have been proposed to model population
growth.
REAL-WORLD PREDICTIONS

An important part of the modeling process,

is to interpret
our mathematical conclusions as real-world
predictions and test them against real data.
Benjamin Gompertz
Benjamin Gompertz was an English self-educated
mathematician and actuary, who is best known for his "law of
mortality".

On 10 October 1810 Gompertz married Abigail Montefiore,

who came from a wealthy Jewish family with strong links with
the stock exchange, at the Hambro Synagogue, London.
Gompertz himself joined the stock exchange in 1810 and he
became a Fellow of the Royal Society in 1819. The following
year he read a paper to the Society which applied the
differential calculus to the calculation of life expectancy.
In 1824, the year his 10 year old son died, Gompertz was
appointed as actuary and head clerk of the Alliance Assurance
Company.
Gompertz applied the calculus to actuarial questions and he is
best remembered for Gompertz's Law of Mortality. Gompertz,
in 1825, showed that the mortality rate increases in a
geometric progression .Hence, when death rates are plotted on
a logarithmic scale, a straight line known as the Gompertz
function is obtained. It is the most informative actuarial
function for investigating the ageing process.
The Gompertz model has been in use as a growth model
even longer than its better known relative, the logistic
model . The model, referred to at the time as the Gompertz
theoretical law of mortality, was first suggested and first
applied by Mr. Benjamin Gompertz in 1825 .He fitted it to
the relationship between increasing death rate and age,
what he referred to as “the average exhaustions of a man’s
power to avoid death”, or the “portion of his remaining
power to oppose destruction”. The insurance industry
quickly started to use his method of projecting death risk.
T h e G o m p e r t z law
Living organisms provide one of the best examples of populations. Often neither
the Malthusian nor logistic models of population growth are appropriate for
biological organisms. One such model assumes that cell populations P(t)
evolve according to the G o m p e r t z equation

With a and b both positive. This model shows a maximum sustainable population of
 A Reaction diffusion system

The reaction diffusion system describes mathematically the process in which the concentrations of

one or more than one substance (usually chemicals) change spatially and temporally. These two

changes are brought about by the diffusion term and the reaction term respectively.

 A Reaction term is the term by which changes in concentration of the reacting substances

occur.

 A Diffusion term is the term by which the substance spreads out or diffuses spatially.

Mathematically reaction diffusion equations can be put succinctly as one equation which usually
 𝜕𝒖 2
= 𝐷 ∇ 𝒖+ 𝒇 ( 𝒖 )
𝜕𝑡

Where

Some appropriate initial and boundary conditions are given, depending on the problem under

consideration.

could be bounded or unbounded. Reaction diffusion equations can be either two-component reaction

diffusion systems or one-component, where is the concentration of the substance under consideration or a population

at time t.

is the Laplacian operator which represents diffusion D being the diffusion coefficient, is the reaction term and

is a smooth function Usually this describes some change like birth or death in some population, or a chemical reaction to give a

few examples
 There are many phenomena exhibiting oscillations in nature. Apart from naturally occurring

oscillations there are the man-made ones in various man-made devices. In the study of physical

systems, one of the earliest concepts introduced is that of the linear, harmonic oscillator [3]. A system

exhibiting harmonic oscillations is usually modelled by more or less similar differential equations. In

addition to the so-called harmonic oscillators, there are the “Nonlinear oscillators”, cropping up in

various branches of the Natural Sciences. Eventually it is realised that “Nonlinear oscillations” are as

essential as harmonic oscillations.

To give some fundamental and varied examples:

 The heart and its stable oscillations

 Josephson junctions or devices generating voltage oscillations of an extremely high frequency;

 Chemical concentrations of substances of a chemical reaction oscillating temporally, for example in

the Belousov–Zhabotinsky reaction.

 The Fisher equation is a one dimensional reaction diffusion equation. That is, the reaction diffusion

system has a single component in the spatial dimension :

where D is a constant, with the initial distribution given in the interval .

This was the work of R. A. Fisher in which he examined the properties of a differential equation

determining the spread of a favoured gene. The population which Fisher studied was one which has

uniform density. When an advantageous mutation occurs in a population this gene will be the favoured

gene.

According to Fisher this advantageous gene then diffuses into the population. Fisher hypothesizes that the

diffusion of the advantageous gene takes the form of an advancing wave. In the Fisher equation this gene

diffusion is compared, in a broad manner, to physical diffusion keeping in mind that diffusion is an

extremely complex process.

The constant in equation is the coefficient related to diffusion, similar to the process of diffusion in

physics. Of course this comparison with diffusion in physics is valid when the distances covered by the

mutating gene are very small. In actuality this comparison of diffusion with physical diffusion is an

oversimplification.
The Fisher Kolmogorov Petrovskii and Piskunov (FKPP) Equation

At this point one should mention the names of Petrovskii and Piskunov along with Fisher and

Kolmogorov. Because of these four names the equation is referred to in some texts as the FKPP equation.

A particular form of a term in the equation

was chosen by these authors as . Such a choice of would give, in the absence of diffusion two
equilibrium points and .
The classic nonlinear reaction diffusion equation due to Fisher is

The term represents a logistic growth term.This can also be understood if one

compares this with the familiar Logistic equation

where it is apparent that the growth of the population is limited by the carrying
capacity of the population normalized to1. is the intrinsic growth rate
The term in the Fisher equation which was referred to as the Logistic term earlier,
models the increase of the frequency of the gene
 The Belousov Zhabotinsky reaction
In the early 1950s a Soviet biochemist, Boris P. Belousov, was trying to
develop a simple chemical model of the oxidation of organic molecules
in living cells.
Central to these pathways is the Krebs cycle, whereby organic acids are
oxidized to In aerobic organisms, oxygen is the oxidizing agent, and the
reactions are catalyzed by enzymes and electron-transport
proteins, many of which rely on iron ions (Fe 2+/ Fe3+) to move electrons
around.
In his test tube version of metabolism, Belousov used citric acid (one of the

intermediates of the Krebs cycle) as an organic substrate, bromate ions () as

oxidizing agent, and cerium ions as catalyst.

Any chemist would expect the reaction to proceed monotonically to equilibrium,

perhaps showing one visible sign of progress by changing from a colorless solution

(cerium in the reduced state, ) to pale yellow (the oxidized state, ).

•So we can imagine Belousov's surprise when his reaction
mixture turned yellow then colorless, then yellow again
and colorless, oscillating dozens of times between
oxidized and reduced states
 Consider chemical equations of the form

𝐴+ 𝐵 → 𝐶 + 𝐷

where represent some chemicals.

By the law of mass action [118], it can be said that “the rate of a reaction either forward or backward is

proportional to the product of the concentration of the reactants”.

here refer to the concentrations of the chemicals and respectively, the rate constant is , the reaction

rate is ,
In the chemical equation (4.2) it is apparent that the change of (concentration of A) with respect to time is

proportional to multiplied with is used up in the reaction and therefore, is taken to be negative. The

related differential equations pertaining to each chemical is written as follows

In general an equilibrium reaction can be represented as

Using the law of mass action the forward reaction can be written as:

(taking into account the number of molecules of each reactants)

Reaction rates

Let us now take a simple reaction

It is apparent that the chemical is consumed twice as quickly as the chemical . To take into account, this

factor, the stoichiometric coefficient of is taken to be

being the rate of the reaction.

We will be investigating the Brusselator, a hypothetical system close in its nature to the BZ reaction
The two species of interest to us are X and Y, the autocatylitic species. The
differential equations given in dimensionless form for these species are:
0
0
Then the system maybe analyzed using usual techniques, to determine its stability and so on.
Balthazar van der Pol (1889-1959) was a Dutch electrical engineer who initiated modern

experimental dynamics in the laboratory during the 1920's and 1930's. He, first, introduced his

(now famous) equation in order to describe oscillations in electrical circuits, in 1927. The

mathematical model for the system is a well known second order ordinary differential equation

with cubic nonlinearity – the Van der Pol equation. Since then thousands of papers have been

published achieving better approximations to the solutions occurring in such non linear systems.

The Van der Pol oscillator is a classical example of self oscillatory system and is now considered

as very useful mathematical model that can be used in much more complicated and modified

systems
But, why this equation is so important to mathematicians, physicists
and engineers and is still being extensively studied?
During the first half of the twentieth century, Balthazar van der Pol pioneered the
fields of radio and telecommunications . In an era when these areas were much
less advanced than they are today, vacuum tubes were used to control the flow of

electricity in the circuitry of transmitters and receivers.

Van der Pol, in 1927, experimented with oscillations in a vacuum tube triode
circuit and concluded that all initial conditions converged to the same periodic
orbit of finite amplitude. Since this behavior is different from the behavior of
solutions of linear equations, van der Pol proposed a nonlinear differential
equation

commonly referred to as the (unforced) van der Pol equation , as a model for the
behavior observed in the experiment.
The Equation is a simple harmonic oscillator with a nonlinear damping

term The damping is positive for and negative for . This results large

amplitude oscillations to decay but when they become too small they are

pumped back up. The energy decreased in one cycle balances the energy pumped in and

the system settles into a self-sustained oscillations.The Equation can be written as the

system of two first order differential equations as

Phase plane of van der Pol equation for
Variations of (left) and (right) with respect to time for
Since its introduction in the 1920’s, the Van der Pol equation has been a
prototype for systems with self-excited limit cycle oscillations. The classical
experimental setup of the system is the oscillator with vacuum triode. The
investigations of the forced Van der Pol oscillator behaviour have carried out by
many researchers. The equation has been studied over wide parameter regimes,
from perturbations of harmonic motion to relaxation oscillations. It was much
attention dedicated to investigations of the peculiarities of the Van der Pol
oscillator behaviour under external periodic (sinusoidal) force and, in particular,
the synchronization phenomena and the dynamical chaos appearing
The Van der Pol equation is now concerned as a basic model for
oscillatory processes in physics, electronics, biology, neurology, sociology and
economics .Van der Pol himself built a number of electronic circuit models of the
human heart to study the range of stability of heart dynamics.
His investigations with adding an external driving signal were analogous to the
situation in which a real heart is driven by a pacemaker. He was interested
infinding out, using his entrainment work, how to stabilize a heart's irregular
beating or "arrhythmias". Since then it has been used by scientists to model a
variety of physical and biological phenomena.
The Hodgkin Huxley Model
By the late 1930s, researchers had come to understand several important things
about the conduction of signals within neurons. For example, they knew that
signaling within neurons is electrical in nature (as opposed to signaling between
neurons, which is usually chemical), and that it occurs in bursts of activity called
action potentials. And they knew that action potentials are stimulated by the
movement of sodium ions across the neuronal membrane through proteins called
ion channels. But the full details of what is going on during an action potential
were not made completely clear until Alan Hodgkin and Andrew Huxley started
collaborating on the issue in 1939.
What is an action potential?
To understand Hodgkin and Huxley's findings, it helps to have some background on
what happens during an action potential. When a neuron is at rest, there are a
variety of charged particles called ions that are unevenly distributed inside and
outside of the cell. Ions are simply atoms that have either gained or lost an
electron; this gain or loss of electrons gives the atoms a negative or positive overall
charge, respectively. When a neuron isn't excited, positively-charged sodium ions
accumulate outside of the neuron, while positively-charged potassium ions
accumulate inside.
There are also negatively-charged chloride ions that accumulate outside and
organic ions that accumulate inside the cell. A number of mechanisms, including
both passive (e.g. diffusion) and active (e.g. the Na-K pump) processes, work to
maintain a unequal balance of positively- and negatively-charged particles between
the inside and outside of the neuron. This difference in charge is known as the
resting membrane potential; typically in neurons it is around -65 to -70 mV, which
means that the inside of the neuron is negatively-charged with respect to the
outside.
One of the difficulties in understanding action potentials before Hodgkin and Huxley's work
was that neurons are incredibly small. (At their largest they are about 100 micrometers, but
they can be under 10 micrometers. By comparison, a human hair is about 80 micrometers.)
Scientists found that the size of the axons in most species made it difficult or impossible to
insert a recording device to measure voltage changes during an action potential.
Hodgkin and Huxley got around this problem by studying action potentials in the relatively
enormous axons (up to 1 mm in diameter) of the squid. They inserted a fine capillary
electrode into the squid giant axon and were able to measure electrical changes within the
axon during an action potential.
 THE HODGKIN-HUXLEY MODEL

In 1952 Hodgkin and Huxley (Hodgkin A. L., 1952), (J.Sneyd, 1998) carried out experiments

to determine how the nerve impulses are transmitted in a neuron. They used the technique o

voltage–clamping on the giant squid axon and collected a large volume of experimental data

They wrote five papers explaining their experimental work and its results.
in the nervous system nerve cells communicate with each other using an electro-chemical mechanism.
Hodgkin and Huxley studied this phenomenon in squids and determined the exact nature of signaling
used by neurons (Hodgkin A. L., 1952). Thus briefly it can be put that these impulsesare exchanged by
the axon through the movement of Sodium (Na+) and Potassium (K+) ions.

A neuron
The excitation and propagation of these impulses arose from the flow of Sodium (Na+) and Potassium

(K+) ions. Thus the model includes a voltage type variable with cubic nonlinearity and a second variable

–“recovery variable” which has linear dynamics. A constant term was added to the cubic representing

injection of an external current or an external stimulus into the nerve membrane. So when that would

represent a rest point or no stimulation. The nature of the stimulation determines how the impulse is

transmitted. So can be a constant or a variable.

The Hodgkin-Huxley system consists of four state
variables. The Hodgkin-Huxley system is
where is membrane potential, is Na (Sodium) activation, is K (Potassium) activation and is Na (Sodium)

inactivation, is the potential, g’s are constant conductances, for example is Sodium conductance which

consists of three activating subunits and one inactivating subunit, and are constant equilibrium

potentials, and are variables bounded by 0 and 1 which are determined by the differential equations
The End