Linear Algebra and Differential Equations

Lecture 24
Lotka-Volterra Equations

Aug 1, 2016

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Introduction

The equations bearing their name were proposed independently in the

1920s by Lotka, who was a chemist studying the interaction of two
chemical compounds, and Volterra who was an applied mathematician
studying ecology.

Volterra was trying to explain the puzzling fish harvest in the Adriatic after
Wold War I. It was assumed that since there had been no fishing during
the war the fish would be abundant but it was found that they were scarce
instead.

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The original Lotka-Volterra equations

Let x denote the density of prey fish and y denote the density of predator
fish. Assume the equations
x 0 = x(a by )
y 0 = y (c + dx)
where a, b, c, d > 0
The idea of the first equation is that in the absence of predators, the prey
would grow at a constant rate a, but decreases linearly as a function of the
density y of the predators. Similarly, in the absence of prey, the density of
predators would decrease but the rate increases proportionally to the
density of the prey.

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The x nullclines are the y axis and the line y = ba . The y nullclines are the
x axis and the line x = dc . The equilibrium points are the points of
intersection of the nullclines i.e. the points
c a
(0, 0)
,
d b
Calculating the Jacobian gives

a by
J=
dy
So


J(0, 0) =

and

a 0
0 c

c a  0
,
= ad
J
d b
b

bc
d
0

bx
c + dx


(0,0) is a saddle


c a
,
is a center
d b

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Phase Plane and trajectories

The trajectories are closed periodic orbits about the non trivial equilibrium
point.

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A more realistic model

The original model is modified by adding a logistic term to each equation
x 0 = x(a ex by )
y 0 = y (c + dx fy )
with a, b, c, d, e, f > 0. The nullclines are the axes and the lines
a ex by = 0

c + dx fy = 0

The first line has negative slope and the second

has a positive slope. The
a
first line intersects the positive x axis at e , 0 . The second line doesnt
intersect
the positive y axis. There are equilibrium points at the origin and
at ea , 0 . There will be a third equilibrium if the two lines intersect.

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Case I : the lines dont intersect

If the lines dont intersect then

c
d

> ea .

Calculating the Jacobian gives


a by 2ex
J=
dy

J(0, 0) =
and

a 0
0 c

bx
c + dx 2fy


(0,0) is a saddle


  a
ba
e
J
,0 =
0 c + da
e
e
a

this has two negative eigenvalues and so it is a stable node.

So ultimately the predators become extinct
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Phase plane plots

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Case II : The lines intersect

In this case

c
d

<

a
e

and the lines intersect at



bc fa da ec
? ?
,
(x , y ) =
bd + ef bd + ef

The Jacobian at (x ? , y ? ) is

J=
tr(J) = ex ? fy ? < 0

ex ? bx ?
dy ? fy ?

det(J) = (bd + ef )x ? y ? > 0

This implies (x ? , y ? ) is stable. (It is a spiral but proving it involves a lot of

algebra).

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Phase plane plots

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Two competing species

In this model each species inhibits the growth of the other as well as
incorporating the logistic style inhibition of other members of the species.
The model equations are
x 0 = x(a bx cy )
y 0 = y (d ex fy )
with a, b, c, d, e, f > 0.
The nullclines both have negative slope and there are four distinct senarios
1

Lines dont intersect and the y nullcline is greater than the x nullcline

Lines dont intersect and the x nullcline is greater than the y nullcline

Lines do intersect and the y nullcline is steeper than the x nullcline

Lines do intersect and the x nullcline is steeper than the y nullcline

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Scenario 1
For the model equations
x 0 = x(1 x y )
y 0 = y (6 2x 3y )
The critical points are (0,0),(0,2),(1,0) and the Jacobian is


1 2x y
x
J=
2y
6 2x 6y
at the critical points


1 0
J(0, 0) =
0 6


J(0, 2) =

1 0
4 6


J(1, 0) =

1 1
0
4

So (0,0) is an unstable node, (0,2) is a stable node, and (1,0) is a saddle.

Eventually x becomes extinct.
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Phase plane plots

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Scenario 2
For the equations
x 0 = x(6 3x 2y )
y 0 = y (1 x y )
The critical points are (0,0),(0,1),(2,0) and the Jacobian is


6 6x 2y
2x
J=
y
1 x 2y
at the critical points


6 0
J(0, 0) =
0 1


J(0, 1) =

4
0
1 1


J(2, 0) =

6 4
0 1

So (0,0) is an unstable node, (0,1) is a saddle, and (2,0) is a stable node.

Eventually y becomes extinct.
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Phase plane plots

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Scenario 3
For the equations
x 0 = x(2 x 2y )
y 0 = y (2 2x y )
The critical points are (0,0),(0,2),(2,0),( 32 , 32 ) and the Jacobian is


2 2x 2y
2x
J=
2y
2 2x 2y
at the critical points



 
  2

2 2
4 2
2 4
3 43
J(0, 2) =
J(2, 0) =
J
,
=
0
1
0 2
43 23
3 3


So (0,0) is an unstable node, 32 . 23 is a saddle, and (2,0) and (0,2) are
stable nodes. Eventually either x or y becomes extinct, depending on the
initial conditions.
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Phase plane plots

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Scenario 4
For the equations
x 0 = x(2 x 2y )
y 0 = y (2 2x y )
The critical points are (0,0),(1,0),(0,1),( 32 , 32 ) and the Jacobian is

J=

2 4x y
y

x
2 x 4y

at the critical points


J(0, 0) =

2 0
0 2


J(1, 0) =

2 1
0
1

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J(0, 1) =

1
0
1 2


J

2 2
,
3 3


=

43
23

32
34

So (0,0) is an unstable node and (2,0)

and (0,2) are saddles nodes.
The characteristic equation at 23 . 23 is
4
8
2 + + = 0
3
3
Thus

2 2
3.3

2
= , 2
3

is a stable node and the system eventually ends up there.

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Phase plane plots

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