Leslie matrix
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The Leslie matrix is a discrete, age-structured model of population growth that is very popular in
population ecology Population ecology is a sub-field of ecology that deals with the dynamics of species populations and how these populations interact with the environment, such as birth and death rates, and by immigration and emigration. The discipline is import ...
named after Patrick H. Leslie. The Leslie matrix (also called the Leslie model) is one of the most well-known ways to describe the growth of populations (and their projected age distribution), in which a population is closed to migration, growing in an unlimited environment, and where only one sex, usually the
female Female ( symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction. A female has larger gametes than a male. Females ...
, is considered. The Leslie matrix is used in
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...
to model the changes in a population of organisms over a period of time. In a Leslie model, the population is divided into groups based on age classes. A similar model which replaces age classes with ontogenetic stages is called a Lefkovitch matrix, whereby individuals can both remain in the same stage class or move on to the next one. At each time step, the population is represented by a
vector Vector most often refers to: *Euclidean vector, a quantity with a magnitude and a direction *Vector (epidemiology), an agent that carries and transmits an infectious pathogen into another living organism Vector may also refer to: Mathematic ...
with an element for each age class where each element indicates the number of individuals currently in that class. The Leslie matrix is a square matrix with the same number of rows and columns as the population vector has elements. The (i,j)th cell in the matrix indicates how many individuals will be in the age class ''i'' at the next time step for each individual in stage ''j''. At each time step, the population vector is multiplied by the Leslie matrix to generate the population vector for the subsequent time step. To build a matrix, the following information must be known from the population: * n_x, the count of individuals (''n'') of each age class ''x'' * s_x, the fraction of individuals that survives from age class ''x'' to age class ''x+1'', * f_x,
fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
, the
per capita ''Per capita'' is a Latin phrase literally meaning "by heads" or "for each head", and idiomatically used to mean "per person". The term is used in a wide variety of social sciences and statistical research contexts, including government statistic ...
average number of female offspring reaching n_0 born from mother of the age class ''x''. More precisely, it can be viewed as the number of offspring produced at the next age class b_ weighted by the probability of reaching the next age class. Therefore, f_x = s_xb_. From the observations that n_0 at time ''t+1'' is simply the sum of all offspring born from the previous time step and that the organisms surviving to time ''t+1'' are the organisms at time ''t'' surviving at probability s_x, one gets n_ = s_xn_x. This implies the following matrix representation: : \begin n_0 \\ n_1 \\ \vdots \\ n_ \\ \end_ = \begin f_0 & f_1 & f_2 & \ldots & f_ & f_ \\ s_0 & 0 & 0 & \ldots & 0 & 0\\ 0 & s_1 & 0 & \ldots & 0 & 0\\ 0 & 0 & s_2 & \ldots & 0 & 0\\ \vdots & \vdots & \vdots & \ddots & \vdots & \vdots\\ 0 & 0 & 0 & \ldots & s_ & 0 \end \begin n_0 \\ n_1 \\ \vdots\\ n_ \end_ where \omega is the maximum age attainable in the population. This can be written as: :\mathbf_ = \mathbf\mathbf_t or: :\mathbf_ = \mathbf^t\mathbf_0 where \mathbf_t is the population vector at time ''t'' and \mathbf is the Leslie matrix. The dominant
eigenvalue In linear algebra, an eigenvector () or characteristic vector of a linear transformation is a nonzero vector that changes at most by a scalar factor when that linear transformation is applied to it. The corresponding eigenvalue, often denoted ...
of \mathbf, denoted \lambda, gives the population's asymptotic growth rate (growth rate at the stable age distribution). The corresponding
eigenvector In linear algebra, an eigenvector () or characteristic vector of a linear transformation is a nonzero vector that changes at most by a scalar factor when that linear transformation is applied to it. The corresponding eigenvalue, often denoted ...
provides the stable age distribution, the proportion of individuals of each age within the population, which remains constant at this point of asymptotic growth barring changes to vital rates. Once the stable age distribution has been reached, a population undergoes
exponential growth Exponential growth is a process that increases quantity over time. It occurs when the instantaneous rate of change (that is, the derivative) of a quantity with respect to time is proportional to the quantity itself. Described as a function, a ...
at rate \lambda. The characteristic polynomial of the matrix is given by the
Euler–Lotka equation In the study of age-structured population growth, probably one of the most important equations is the Euler–Lotka equation. Based on the age demographic of females in the population and female births (since in many cases it is the females that are ...
. The Leslie model is very similar to a discrete-time Markov chain. The main difference is that in a Markov model, one would have f_x+s_x=1 for each x, while the Leslie model may have these sums greater or less than 1.


Stable age structure

This age-structured growth model suggests a steady-state, or stable, age-structure and growth rate. Regardless of the initial population size, N_0, or age distribution, the population tends asymptotically to this age-structure and growth rate. It also returns to this state following perturbation. The Euler–Lotka equation provides a means of identifying the intrinsic growth rate. The stable age-structure is determined both by the growth rate and the survival function (i.e. the Leslie matrix).Further details on the rate and form of convergence to the stable age-structure are provided in Charlesworth, B. (1980) Evolution in age-structured population. Cambridge.
Cambridge University Press Cambridge University Press is the university press of the University of Cambridge. Granted letters patent by King Henry VIII in 1534, it is the oldest university press in the world. It is also the King's Printer. Cambridge University Pre ...
For example, a population with a large intrinsic growth rate will have a disproportionately “young” age-structure. A population with high mortality rates at all ages (i.e. low survival) will have a similar age-structure.


Random Leslie model

There is a generalization of the population growth rate to when a Leslie matrix has random elements which may be correlated.M.O. Caceres and I. Caceres-Saez, Random Leslie matrices in population dynamics, J. Math. Biol. (2011) 63:519–556 DOI 10.1007/s00285-010-0378-0 When characterizing the disorder, or uncertainties, in vital parameters; a perturbative formalism has to be used to deal with linear non-negative
random matrix In probability theory and mathematical physics, a random matrix is a matrix-valued random variable—that is, a matrix in which some or all elements are random variables. Many important properties of physical systems can be represented mathemat ...
difference equations In mathematics, a recurrence relation is an equation according to which the nth term of a sequence of numbers is equal to some combination of the previous terms. Often, only k previous terms of the sequence appear in the equation, for a parameter ...
. Then the non-trivial, effective eigenvalue which defines the long-term asymptotic dynamics of the mean-value population state vector can be presented as the effective growth rate. This eigenvalue and the associated mean-value invariant state vector can be calculated from the smallest positive root of a secular polynomial and the residue of the mean-valued Green function. Exact and perturbative results can thusly be analyzed for several models of disorder.


References


Further reading

* * *{{cite book , last=Pollard , first=J. H. , year=1973 , title=Mathematical models for the growth of human populations , location= , publisher=Cambridge University Press , isbn=0-521-20111-X , chapter=The deterministic theory of H. Bernardelli, P. H. Leslie and E. G. Lewis , pages=37–59 Population Population ecology Matrices