Matris de Leslie

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Matris de Leslie

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This worksheet requires some knowledge of Linear Algebra. The material needed for this

worksheet is covered in Sections 9.2 and 9.3 of Neuhauser (2004) Calculus for Biology

and Medicine. 2nd edition. Prentice Hall. A summary is provided below.

Matrix Algebra

A m �n matrix A = ��aij �

�, i = 1, 2,K , m, j = 1, 2,K n , is a rectangular array of numbers

with m rows and n columns. We write this as

�

a a22 K a2 n �

A = �21 �

�K K K K �

� �

am1

� am 2 K amn �

A 1�n matrix is called row vector, and a m �1 matrix is called a column vector. Two

m �n matrices A and B are the same if all their entries agree. (We can only compare

matrices of the same size.)

Matrices of the same size can be added. Suppose that A and B are m �n matrices. Then

C = A+ B

for i = 1, 2,..., m and j = 1, 2,..., n . Matrix addition satisfies the following two properties:

A+ B = B+ A

( A + B) + C = A + ( B + C )

Matrices in MATLAB

>> u=2:2:10

Worksheet 5: Age-structured models—the Leslie matrix

>>v=u.^2

Note the dot before the “^”. It indicates that each component is squared.

To enter a matrix, use semicolons. Entries within the same row may be separated by

commas:

>>A=[1, 0, 3; 2, 4, 5]

To add two matrices, first define two matrices that are of the same size, for instance

>>A=[1,2,3;4,5,6]

>>B=[-1,2,-3;-4,5,-6]

>>C=A+B

The transpose of a matrix A, denoted by A’, is obtained by switching rows and columns.

If you type

>>D=A’

Matrix Multiplication

Matrix multiplication is quite a bit more involved. We give the definition first. Suppose

A=� � �is a m �l matrix and B = �

aij � bij �

� �is a l �n matrix, then

C = AB

is a m �n matrix with

l

cij = ai1b1 j + ai 2b2 j + ... + ail blj = �aik bkj

k =1

for 1 �i �m and 1 �j �n . Note that in this product, the number of columns of the matrix

A must be the same as the number of rows of the matrix B.

2

Worksheet 5: Age-structured models—the Leslie matrix

�1 2 3 -3�

�1 2 3 � � �

A=� � and B = �0 -1 4 0 �

�-1 0 4 �

�

�- 1 0 -2 1 �

�

Since the number of columns of A is the same as the number of rows of B, the product

C = AB is defined, and C is a 2 �4 matrix. To find the entry in say the second row and

the third column of the matrix C, you would multiply the numbers of the second row of

matrix A by the numbers of the third column of the matrix B.

�3 �

c23 = [ -1 0 4] � �

�4 �= (-1)(3) + (0)(4) + (4)( -2) = -3 + 0 - 8 = -11

�

�-2 �

�

>>A=[1,2,3;-1,0,4]

>>B=[1,2,3,-3;0,-1,4,0;-1,0,-2,1]

>>C=A*B

enter the following matrix

>>D=[1 2;3 4]

>>D*D

>>D^2

Make sure you distinguish this from the following two operations that use a dot before

the operator. The first one multiplies each of the corresponding entries (and is thus

different from matrix multiplication):

>>D.*D

Likewise,

>>D.^2

takes the square of each entry but does not square the matrix.

3

Worksheet 5: Age-structured models—the Leslie matrix

spend 13-17 years in the nymphal stage; they only reproduce once in their lifetime. Many

animals, such as humans, elephants, etc., do not reproduce during their first years and

then their reproductive success is age-dependent. To model such situations, age-

dependent population models are appropriate. Patrick Leslie introduced matrix models

that have discrete age classes with synchronous reproduction (Leslie 1945). The models

are parameterized by age-specific survival probabilities and average number of female

offspring.

An age-structured population with k age classes is described by a population

vector of length k

n1 (t )

n (t )

n(t ) =

2

nk (t )

where n j (t ) denotes the number of females in age class j. Individuals within an age

class are assumed to have equal birth and death probabilities. Survival from age class j to

j+1 is given by probability Pj; female individuals in age class j give birth on average to Fj

female offspring. This can be illustrated using life cycle graphs (see Figure 1):

F4

F3

F2

P1 P2 P3

F1 1 2 3 4

4

Worksheet 5: Age-structured models—the Leslie matrix

n1 (t + 1) = F1n1 (t ) + F2 n2 (t ) + + Fk n k (t )

n2 (t + 1) = P1n1 (t )

n3 (t + 1) = P2 n 2 (t )

nk (t + 1) = Pk -1n k -1 (t )

n1 (t + 1) F1 F2 Fk n1 (t )

n (t + 1) P 0 0 n2 (t )

2 = 1

0 P2 0

nk (t + 1) 0 0 Pk -1 0 nk (t )

The matrix contains both the survival probabilities (just below the diagonal) and the

fecundities (in the first row). The matrix is called Leslie matrix. Take a moment to verify

that the two descriptions are equivalent.

Task 1:

0 2.3 1.4

L=

0.4 0 0

0 0.6 0

(a) Translate the matrix into a life cycle graph. (b) If at time 0, there are 100 individuals in

age class 1, and no individuals in each of age classes 2 and 3, how many individuals are in the

three age classes at time 2? You may use MATLAB to solve this. Note that if you enter the

population vector at time 0 as a row vector, you need to transpose that vector first before

you can compute the population vector at time 1.

To analyze matrix models, we need to learn some tools from linear algebra.

Definition: Assume that A is a square matrix. A nonzero vector w that satisfies the

equation

Aw = w

is a (right) eigenvector of the matrix A and the number λ is the corresponding eigenvalue.

Note that we require that the eigenvector is different from the zero vector. An eigenvalue

can be 0, it can even be a complex number. The geometric interpretation of eigenvectors

5

Worksheet 5: Age-structured models—the Leslie matrix

gets mapped by the matrix on the line through the eigenvector. The eigenvalue allows us

to find the length of the mapped vector.

To find eigenvectors and eigenvalues, we need to solve the equation

Aw = w

Aw - w = 0

( A - I ) w = 0

where I is the identity matrix. (The identity matrix is a square matrix that has 1s on the

diagonal line and 0s elsewhere.) In order to obtain a nontrivial solution (i.e., w 0 ), the

matrix A - I must be singular. The determinant of a matrix can be used to check

whether a matrix is singular, and the condition is:

det( A - I ) = 0

Solving this equation will yield values for λ. For each value of λ, we can then find an

eigenvector w. Determinants were originally introduced to determine whether a system of

linear equations has a unique solution. The theory of determinants has moved far beyond

this. But to us, this will be the situation where we will encounter determinants.

If A is a two-by-two matrix, the computation can be carried out by hand. Namely,

if

a a12

A = 11

a 21 a 22

then

a - a12

det( A - I ) = det 11 = a11 - a 22 - - a12 a 22

a 21 a 22 -

Multiplying this out yields a quadratic equation, which can be solved using

- b b 2 - 4ac

ax 2 + bx + c = 0 has solution x1, 2 =

2a

If all eigenvalues of the matrix A are distinct and real, the eigenvectors will be real and

can be computed by solving Aw = w for each eigenvalue.

Determinants in MATLAB

6

Worksheet 5: Age-structured models—the Leslie matrix

Then the command is

>>det(A)

To use MATLAB to find the eigenvalues of a matrix using the determinant, the following

commands are used (below, we’ll introduce a quicker way):

>>syms lam

>>A=[2 3;1 4];

>>I=[1 0;0 1];

>>solve(det(A-lam*I));

1 2�

�

A=� �

3 2�

�

than the series of commands we used previously:

>>eig(A)

Factoring the right-hand side and then setting it equal to zero yields

( + 1)( - 4) = 0

and hence the same answer.

The eigenvectors can be computed by hand once the eigenvalues are known by

solving the equation

Au = u

In the case of the eigenvalue 1 = -1 , this amounts to solving

u1 �

1 2 ��

� �u�

� �� �= (-1) �1 �

u2 �

3 2 ��

� u2 �

�

7

Worksheet 5: Age-structured models—the Leslie matrix

u1 + 2u2 = -u1

3u1 + 2u2 = -u2

This reduces to just one equation since the two equations are identical:

u1 + u2 = 0

�1 �

This has infinitely many solutions. We need to pick one, for instance, the vector � �

�-1�

satisfies the equation.

This produces two matrices. The matrix D is a diagonal matrix with the eigenvalues along

the main diagonal. The columns of the matrix V are the eigenvectors scaled to length 1.

In this case, we find

V =� � and D = � �

�0.7071 0.8321 � �0 4 �

In Step 2b of the Computer Lab, we will compute the fraction of individuals in each of

the age classes. If these fractions stabilize over time, they represent the stable age

distribution. A stable age distribution does not always exist. Suppose that

w1 = w11 , w12 ,..., w1k is a stable age distribution, then applying the Leslie matrix to this

vector, that is computing Lw1 , results in a new vector that can be scaled so that it is equal

to w1 . This means that w1 is a right eigenvector. The scaling can be expressed as

Lw1 = 1w1 , implying that 1 is the corresponding eigenvalues.

We will now use an important result from linear algebra, which we cannot prove here.

Assume that the eigenvalues of the Leslie matrix are real and distinct. Because of the

assumption of distinct eigenvalues, the corresponding right eigenvectors are also distinct,

and we can write any initial population vector n(0) as a linear combination of the right

eigenvectors of the Leslie matrix

k

n ( 0) = ci wi

i =1

8

Worksheet 5: Age-structured models—the Leslie matrix

where the coefficients ci are determined by the initial vector n(0). Applying the Leslie

matrix to this, we see what the advantage of writing the initial vector in this way is

k k k

n(1) = Ln (0) = L ci wi = ci Lwi = ci i wi

i =1 i =1 i =1

k k k

n(2) = Ln(1) = L ci i wi = ci i Lwi = ci i2 wi

i =1 i =1 i =1

k k k

n(t ) = Ln (t - 1) = L ci ti -1 wi = ci ti -1 Lwi = ci ti wi

i =1 i =1 i =1

We find that the population vector at time t is a weighted average of exponentials ti

where the weights are determined by the initial population vector.

If all the eigenvalues are distinct, we can order them so that 1 2 3 . We then find

that for t large, the largest eigenvalue dominates in the sense that

t t t

k

n(t ) = 1t ci i wi =1t c1 w1 + c 2 2 w2 + + c k k w2

1 1

i =1

1

0 as t

n (t )

lim = c1 w1

1t

t

The eigenvector w1 corresponding to the largest eigenvalue 1 is also the stable age

distribution. The largest eigenvalues 1 tells us how quickly the population grows for

large times. We see that when 1 1 , the population will grow exponentially without

bound, and when 1 < 1 , it will decline exponentially fast.

Some Generalizations

So far, we assumed that all eigenvectors are distinct. We wish to relate this property to

biological properties of the population. We begin with a number of definitions.

9

Worksheet 5: Age-structured models—the Leslie matrix

A matrix is nonnegative if all its entries are nonnegative. A matrix is positive if all its

entries are positive. Note that all Leslie matrices are nonnegative.

Irreducibility

A nonnegative matrix or a life cycle graph is irreducible if and only if every stage can be

reached from every other stage. In terms of the life cycle graph this means that there is a

path from every node to every other node.

Life cycle graphs with postreproductive age classes do not result in an irreducible matrix.

Primitivity

positive.

A loop is a path that has the same start and end point and passes through each node at

most once.

A life cycle graph is primitive if it is irreducible and the greatest common divisor of the

lengths of its loops is 1 (see Figure 2).

Figure 2: The life cycle graph on the left is not primitive since there is only one loop and it is of length 3.

The life cycle graph on the right is primitive since there are two loops, one of length 2, the other of length

3; consequently, the greatest common divisor is 1.

primitive.

Case 1

n(t ) = c w

i

t

i i with 1 j , j 1

In this case,

10

Worksheet 5: Age-structured models—the Leslie matrix

n (t )

lim = c1 w1

t 1t

and the right eigenvector w1 corresponding to the largest eigenvalue is a stable age

distribution (see Computer Lab, Steps 1-2).

Case 2

If the Leslie matrix has postreproductive classes, it is reducible. The Leslie matrix can be

reduced into four submatrices

B 0

L= 1

B21 B2

The matrix B1 alone gives the growth rate and is unaffected by the post reproductive age

classes (see Computer Lab, Step 3).

Case 3

1 0

n(t ) = ci wi with

t

i 1 = j , j = 2, , d

, jd

1 j

11

Worksheet 5: Age-structured models—the Leslie matrix

Read the steps below.

Read the article by R.N. Holdway and C. Jacomb. 2000. Rapid Extinction of the

Moas (Aves: Dinornithiformes): Models, Test, and Implications.

Step 1

In the accompanying EXCEL spreadsheet (Tab: “Leslie Matrix”), the following Leslie

Matrix is entered

�

�0.8 0 0 0�

L= � �

�0 0.5 0 0�

� �

�0 0 0.3 0 �

We will run the population for 50 time steps when initially there are 200 individuals in

the first age class and no other individuals. Note that row 9 is different from the other

rows (the other rows are explained at the end of this worksheet). In row 9, we entered the

following in the respective cells:

Cell C9: =G4

Cell D9: =G5

Cell E9: =G6

Cell F9: =SUM(B9:E9)

Cell G9: =F10/F9

Cell H9: =B9/F9

Cell I9: =C9/F9

Cell J9: =D9/F9

Cell K9: =E9/F9

Row 10 is explained at the end of this worksheet. You can drag the rows down so that you

can investigate the behavior for the first 50 time steps.

Step 2

(a) Plot the population sizes of each age class as a function of time for t=0,1,2,…,20. Log

transform the vertical axis and replot. Explain why for sufficiently large t the data points

lie along a straight line. Plot the proportion of each age class as a function of time and use

12

Worksheet 5: Age-structured models—the Leslie matrix

it to find a stable age distribution. (Note that once you transform the vertical axis to a

logarithmic scale, you might get a message from Microsoft Excel telling you that

“Negative or zero values cannot be plotted correctly on log charts….” Simply change the

scale back to the arithmetic scale and you get rid of these messages.)

(b) Column G lists the ratio of consecutive population sizes. What is the significance of

this value? How is it related to the largest eigenvalues?

(c) Columns H-K list the fraction of individuals in each age class. Plot the fraction for

each age class as a function of time for t=0,1,2,…,20. Is there a stable age distribution

and if yes, what is it?

Step 3

Change the matrix from Step 1 so that there are two post reproductive age classes:

1.6 1.5 0 0

0.8 0 0 0

L=

0 0.5 0 0

0 0 P3 0

Check that the growth parameter of the population is unaffected by changes in the

survival probabilities of the post reproductive age classes by increasing P3 from 0 to 1 in

increments of 0.1. Assume that initially there are 200 individuals in the first age class and

no other individuals.

Step 4

0 0 0 5

0.7 0 0 0

L=

0 0.7 0 0

0 0 0.7 0

Assume that initially there are 200 individuals in the first age class and no other

individuals. Describe what you see. Change the survival probabilities from 0.7 to 0.5.

What happens?

Step 5

Steps 1-4 is done more easily in Matlab. Here is the code for Steps 1 and 2:

clear

L=[1.6 1.5 0.25 0.1; 0.8 0 0 0; 0 0.5 0 0; 0 0 0.3 0];

X0row=[200 0 0 0];

13

Worksheet 5: Age-structured models—the Leslie matrix

X0=X0row';

Time=101;

NPop=zeros(4,Time);

NPop(:,1)=X0;

for i=1:Time-1

NPop(:,i+1)=L*NPop(:,i);

end

Gen=0:Time-1;

semilogy(Gen,NPop); xlabel('Generation'); ylabel('Population

Size');title('Leslie Model');

legend('Age 1','Age 2','Age 3','Age 4')

[V,D]=eig(L);

norm=sum(V(:,1));

age=V(:,1)./norm;

Step 6

Comment on the appropriateness of use of Leslie matrices in the article by Holdway and

Jacomb in light of the fact that the Leslie matrix corresponds to exponential

growth/decay.

Literature Cited

P.H. Leslie. 1945. On the Use of Matrices in Certain Population Mathematics. Biometrika

33:183-212.

14

Created by Claudia Neuhauser

Worksheet 5: Age-structured models—the Leslie matrix

A B C D E F G H I J K

1 Leslie Matrix

2 1 2 3 4 Initial vector

3 1 1.6 1.5 0.25 0.1 N1= 200

4 2 0.8 0 0 0 N2= 0

5 3 0 0.5 0 0 N3= 0

6 4 0 0 0.3 0 N4= 0

7

8 Year Stage 1 Stage 2 Stage 3 Stage 4 Total λ_t w_11 w_12 w_13 w_14

9 0 200 0 0 0 200 2.4 1 0 0 0

10 1 320 160 0 0 480 2.266667 0.666667 0.333333 0 0

11 2 752 256 80 0 1088 2.169853 0.691176 0.235294 0.073529 0

12 3 1607.2 601.6 128 24 2360.8 2.174382 0.680786 0.254829 0.054219 0.010166

13 4 3508.32 1285.76 300.8 38.4 5133.28 2.174198 0.683446 0.250475 0.058598 0.007481

Cell C10: =$B$4*B9+$C$4*C9+$D$4*D9+$E$4*E9

Cell D10: =$B$5*B9+$C$5*C9+$D$5*D9+$E$5*E9

Cell E10: =$B$6*B9+$C$6*C9+$D$6*D9+$E$6*E9

Cell F10: =SUM(B10:E10)

Cell G10: =F11/F10

Cell H10: =B10/F10

Cell I10: =C10/F10

Cell J10: =D10/F10

Cell K10: =E10/F10

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