18 Oct 2016

Introtopopulationmatrixmodels
Reviewofmatrixalgebra
Ageclassifiedmatrixmodel:Lesliematrix
Eigenvalueandeigenvectors
SensitivityandElasticity

Stageclassifiedmatrixmodel

POPULATIONMATRIXMODELS

Whymatrixmodels?
Exponential/logisticmodelstreated

individualsinapopulationasallidenticaland
uniform nottrue!
Populationprocesses(e.g.,survival,growth
andreproduction)arenotuniformacrossall
individuals
Onewayofincludingindividualvariabilityis
throughstructuring(e.g.,age,size)
Wanttolooknotonlyatsize,butalso
shape/structureofthepopulation

Applications

Definitions

Matrix - rectangular array of symbols (numbers,

variables, functions, etc.)
Entry or Element of the matrix one of the symbols
it contains

Identified by subscripts denoting row and column

E.g., a32 element in row 3, column 2 of matrix A

Dimensions number of its rows and columns (m x

n)
Column vector m x 1 matrix
Row vector 1 x n matrix
Scalar ordinary number (1 x 1)

18 Oct 2016

Operations: Transpose
4 x1 3 x2 2 x3 0
2 x1 2 x2 5 x3 6
x1 x2 3 x3 1

Transpose of A AT
Switching rows and columns

1 4
T
1 2 3

4 5 6 2 5

3 6

2 x1 0
4 3
2 2 5 x 6

2
1 1 3 x3 1

Operations: Addition

Operations: Multiplication

Add corresponding elements

a11 a12 b11 b12 a11 b11 a12 b12

a
b
a b

a
b
22
21 22 21 21 a22 b22
21

A and B of the same dimensions

A+B=B+A

Scalar multiplication

Multiply each entry in the matrix by the scalar

a b 2a 2b
2
2c 2d
c
d

18 Oct 2016

Operations: Multiplication

Review of matrix manipulations

Matrix multiplication: take the element-wise product

and add them together
b1
a1 a2 a3 b2 a1b1 a2b2 a3b3
b3
# of elements in a row of first matrix has to be the
same as the # of elements in column in second
matrix
A
1x3

A
2x2

B
2x2

a11 a12 b11 b12 a11b11 a12b21 a11b12 a12b22

a
b
a b a b
a
b
22 21
22
21 11 22 21 a21b12 a22b22
21

B
3x1
AB BA

inner coordinates
agree
dimension of
product

TheLeslieMatrix(ageclassifiedmatrix)
Ages 0 1 :
Ages 2 :
Ages 2 3 :

Age class 1
Age class 2
Age class 3

Age is divided into age classes

Population size/state - ni (t)

Given by the numbers of individuals in each age
class.
Individuals in age classes 2 and 3 at time t + 1 are
Survivors of the previous age class at time t

n2 (t 1) P1n1 (t )
n3 (t 1) P2 n2 (t )

Pi = probability an individual of age class i survives

for one time interval (the projection interval)

New members of age class 1 from reproduction

Fi = per capita fertility of age class i

n1 (t 1) F1n1 (t ) F2 n2 (t ) F3 n3 (t )

n1
F1
n (t 1) P
2
1
n3
0

F2
0
P2

F3 n1
0 n2 (t )
0 n3

n(t+1) = An(t)
vector n describes the state of the population
matrix A population projection matrix
(Leslie matrix)
First row fertilities (Fi )
Sub-diagonal survival probabilities (Pi )

18 Oct 2016

Projection

Hughes 1984

Projection

n(1) = A n(0)
time
n(2) = A n(1)
n(3) = A n(2)
n(4) = A n(3)
n(5) = A n(4)
n(6) = A n(5)

Inageclassifiedmatrix,assumethat

variabilityinsurvivalandfecunditylieininter
individualagedifferences

n(t) = At n(0)

Projectpopulationfromtimettotimet+1;

unitoftime=projectioninterval=ageclass
width=important
IfA isconstant=assumeconstant

probabilities

18 Oct 2016

Analyzingthematrixmodel:
EigenvectorsandEigenvalues

Oncestableage/stagedistributionhasbeenattained,theonlyeffectof
multiplyingtheprojectionmatrixbythecolumnvector,N,istoincreasethe
populationsizebythegeometricgrowthfactor(R)

Theeigenvector isavectorsuchthatwhenyoumultiplyitbyamatrixmerely
lengthensorshrinksit(changesonlythemagnitude notdirection)

And,thefactorbywhichitislengthenedorshrunkistheeigenvalue (
scalarvalue);alsocalledthedeterminantofthematrix

AnXbyXmatrixcanhaveuptoXdifferenteigenvectorswithcorresponding
eigenvalues

Mostimportantistheonewiththebiggesteigenvalue(strongestinfluence)

Givenaconstantprojectionmatrix,any

initialagedistributionvectorwillconvergeto
thesamegrowthrateandstableage
distribution theproportionsremain
consistent(notnumbers)
Whenapopulationreachesthestableage

distribution,itgrowsexponentially

EigenvectorsandEigenvalues:
Biologicallyspeaking
Rightdominanteigenvector

stableage/stagedistribution,(w1)
1w1 Aw1
Leftdominanteigenvector
1 v1 v1A
reproductivevalues,(v1)
(contributionofeachstagetothe
populationgrowth)
Dominant(largest)eigenvalue,1
correspondingtotherightdominant
eigenvector geometricgrowth
factor,R;populationgrowthrate

N (t 1) 1N (t )

Matlab: mat1project

18 Oct 2016

Whattheeigenvaluecantellyou
>1:exponentialgrowth
<1:exponentialdecay

Sensitivity & Elasticity Matrices

To determine how the effect of changes in the transitions/processes affect the
population dynamics, particularly the magnitude of the dominant eigenvalue
(growth rate)

Sensitivity: absolute rate of change of 1 with

respect to absolute change in a matrix element

1< <0:dampedoscillationswithperiod=2
Sij

< 1:divergingoscillationswithperiod=2
Complex :oscillations

Dampingratio:largesteigenvalue/second

1
aij

Problem with sensitivity matrix is that you have different value

ranges for variables (e.g., survival: 0 1, fecundities can be
large)... So use standardized sensitivity -> Elasticity

largestvalue howfastthepopulationis
approachingthestablestagedistribution

Sensitivity and Elasticity

Elasticity: relative rate of change of 1 with

respect to relative change in a matrix element
Elasticity analysis estimates the effect of a proportional change
in the vital rates on population growth. In essence, elasticities
are proportional sensitivities, scaled so that they are
dimensionless. This allows you to directly compare survival and
reproductive matrix elements.

1
Eij

1
aij
aij

aij 1 aij
Sij
1 aij 1

matlab:
mat2valvecspitt (senstuff)

18 Oct 2016

Stageclassifiedmatrix
Problemswithusingageclassifiedmatrix:
Longlivedorganisms
Organismsthatcannotbeaged

Assumethatvariabilityinsurvivorshipand

fecundityisduesolelytothecategorization
used

Probabilityof

growing/shrinking
toothersizeclass
subdiagonals
Probabilityof
stayinginsame
sizeclass
diagonal
Firstrow
fecundities

Hughes 1984 Population dynamics

based on individual size rather than age:
a general model with a reef coral
example. The American Naturalist.