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Structured Population Dynamics in a Real World Context
Ana Carolina Loureiro Martins
Thesis to obtain the Master of Science Degree in
Mathematics and Applications
Supervisor: Professor Henrique Manuel dos Santos Silveira de Oliveira
Examination Committee
Chairperson: Professor Maria Cristina de Sales Viana Serodio SernadasMembers: Professor Joao Maria da Cruz Teixeira Pinto
Professor Luıs Humberto Viseu MeloSupervisor: Professor Henrique Manuel dos Santos Silveira de Oliveira
July 2017
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Acknowledgments
After these five tough years, it was a pleasure to make part of this academical community in which
I found what excellence and resilience truly are. I thank every given opportunity to meet new people,
faculty and colleagues, and to enrich myself with new ways of thinking.
I want to specially thank the invaluable and helpful support of my supervisor, Professor Henrique
Oliveira, which gave me the opportunity of working in my thesis while contributing and applying my newly
acquired knowledge in the project called Conhecer Arroios, promoted by Instituto Superior Tecnico de
Lisboa and Junta de Freguesia de Arroios.
Furthermore, I couldn’t have finished this degree without the constant support and hope of my par-
ents, Ana and Joao, and of someone very close to me, Marılia. I also want to express my gratitude to
Miguel Pereira, that read everything carefully to avoid any grammar and spelling mistakes and helped
me finish writing this thesis in the computer when I couldn’t, due to a shoulder surgery.
Last but not least, I want to also thank all of my friends, from LMAC and MMA, that went with me on
this journey.
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Resumo
O estudo de como decorre a evolucao de uma populacao ao longo do tempo e de quao vulneravel
esta se encontra face a mudancas nos parametros que a descrevem e essencial para a compreensao
das necessidades dos habitantes e para a tomada de decisoes segundo um plano a longo prazo. Por-
tanto, a exploracao de ferramentas que permitem esta visao global dos habitantes de uma determinada
regiao e crucial para o desenvolvimento dos mesmos. Nesta tese, e feita uma pequena pesquisa so-
bre modelos populacionais, seguindo-se a apresentacao de novos modelos que contemplam imigracao
e emigracao. Alem disso, o conceito original de matrizes de Leslie e adaptado de forma a integrar
os fluxos migratorios. Uma deducao das caracterısticas da entropia evolucionaria e desenvolvida, e
a entropia da taxa de crescimento e analisada. Um algoritmo que comprime as matrizes de Leslie e
tambem explicado. Finalmente, este conhecimento e aplicado num contexto do mundo real, permitindo
a obtencao de projeccoes populacionais e medidas intrınsecas do ecossistema em estudo.
Palavras-chave: Modelos Populacionais, Matrizes de Leslie, Entropia Evolucionaria, Aplicacao
a um Contexto da Vida Real, Compressao de Matrizes de Leslie.
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Abstract
Understanding how a population evolves with time and how vulnerable to changes in certain param-
eters this ecosystem is is essential to truly comprehend the needs and make decisions according to
long term plans. Therefore, the exploration of tools that help this overview of the inhabitants of a certain
area is crucial to their development. In this thesis, a brief survey about population models is made,
being followed by the presentation of new models that include immigration and emigration. Also, the
original concept of Leslie matrix is adapted in order to integrate migration flows. A deduction of the char-
acteristics of the evolutionary entropy is developed, and the sensitivity of the growth rate is analysed.
An algorithm that compresses Leslie matrices is further explained. Finally, this knowledge is applied
in a real world context, allowing the obtention of population projections and intrinsic measures of the
ecosystem in study.
Keywords: Population Models, Leslie Matrices, Evolutionary Entropy, Application to Real World
Context, Compression of Leslie Matrices.
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Contents
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii
Resumo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
List of Tables . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
List of Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii
1 Introduction 1
1.1 Motivation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Claim of Contributions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.3 Thesis Outline . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Background 3
2.1 Concepts and relevant theorems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.1.1 Matrices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.1.2 Graphs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.1.3 Population matrices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.1.4 Relation between Leslie Matrices and Graphs . . . . . . . . . . . . . . . . . . . . . 7
2.1.5 Population Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2.2 State of the art . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.2.1 Model proposed by H. Leslie [11] . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.2.2 Model proposed by L.P. Lefkovitch [13] . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.2.3 Model proposed by H. Caswell and N. Sanchez Gassen [23] . . . . . . . . . . . . 11
2.2.4 Population Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3 Proposed Models 13
3.1 Model without migratory flows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.2 Model with one migratory flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
3.3 Model with immigration and correction flows . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3.4 Model with internal and external flows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3.5 Model with Leslie matrix that includes immigration and emigration I . . . . . . . . . . . . . 15
3.6 Model with Leslie matrix that includes immigration and emigration II . . . . . . . . . . . . 16
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4 Partial Derivates of Evolutionary Entropy 19
4.1 Evolutionary Entropy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
4.1.1 Characteristics of the Evolutionary Entropy . . . . . . . . . . . . . . . . . . . . . . 20
4.2 Sensitivity Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
5 Compression of the Leslie matrices 25
5.1 Re-weighting the edges of the graph G2c . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
5.2 Proposed Algorithm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
6 Real Word Context 33
6.1 Construction of Arroios’ Leslie Matrices based on 2001 and 2011 data . . . . . . . . . . . 33
6.1.1 Fertility Indices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
6.1.2 Mortality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
6.1.3 Survival Indices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
6.1.4 Immigration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
6.1.5 Emigration to Foreign Countries . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
6.1.6 Error Estimate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
6.2 Application of the models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
6.2.1 Model without migratory flows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
6.2.2 Model with one migratory flow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
6.2.3 Model with immigration and correction flows . . . . . . . . . . . . . . . . . . . . . . 44
6.2.4 Model with internal and external flows . . . . . . . . . . . . . . . . . . . . . . . . . 47
6.2.5 Model with Leslie matrix that includes immigration and emigration I . . . . . . . . . 49
6.2.6 Model with Leslie matrix that includes immigration and emigration II . . . . . . . . 52
6.3 Partial Derivatives of Evolutionary Entropy . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
6.3.1 Leslie matrix without migrations 6.2.1 . . . . . . . . . . . . . . . . . . . . . . . . . 55
6.3.2 Leslie matrix that includes immigration and emigration I 6.2.5 . . . . . . . . . . . . 56
6.3.3 Leslie matrix that includes immigration and emigration II 6.2.6 . . . . . . . . . . . . 56
6.4 Population Sample from 2017 and the Application of Models . . . . . . . . . . . . . . . . . 57
7 Conclusions 61
Bibliography 63
A Iterative Method 65
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List of Tables
5.1 Table with the number of children born, of women that survived from age-class Ci to Ci+1,
and of women by c age-classes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
5.2 Table with the number of children born, of women that survived from age-class Ci to Ci+1,
and of women by 2c age-classes, related with the women subdivided by c age-classes. . . 30
6.1 Fertility rates obtained from 2001’s data without considering migratory flows. . . . . . . . 38
6.2 Survival rates obtained from 2001’s data without considering migratory flows. . . . . . . . 39
6.3 Proportion of the flux relatively to the estimated female residents in 2011 of the model
with one migratory flow in Arroios obtained using 2001 and 2011 Census data. . . . . . . 42
6.4 Proportion of the immigration flow relatively to the estimated female residents in 2011 of
the model with immigration and correction flows in Arroios obtained using 2001 and 2011
Census data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
6.5 Proportion of a correction flow that includes emigration and errors in estimation of immi-
gration relatively to the estimated female residents in 2011 of the model with immigration
and correction flows in Arroios obtained using 2001 and 2011 Census data. . . . . . . . . 45
6.6 Proportion of the internal flow relatively to the total female residents of the model with
internal and external flows in Arroios obtained using 2001 and 2011 Census data. . . . . 47
6.7 Proportion of the external flow relatively to the total female residents of the model with
internal and external flows in Arroios obtained using 2001 and 2011 Census data. . . . . 47
6.8 Fertility rates using the model with Leslie matrix that includes immigration and emigration
II and an iterative method. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
6.9 Survival rates using the model with Leslie matrix that includes immigration and emigration
II and an iterative method. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
6.10 Fertility rates using the model with Leslie matrix that includes immigration and emigration
II and an iterative method. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
6.11 Survival rates using the model with Leslie matrix that includes immigration and emigration
II and an iterative method. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
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List of Figures
4.1 S = 0, where S is a measure of the number of replicative cycles. . . . . . . . . . . . . . . 21
4.2 S > 0, where S is a measure of the number of replicative cycles. . . . . . . . . . . . . . . 21
6.1 Distribution of female newborns using the non-adjusted number and the adjusted number
of female newborns, using 2001 data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
6.2 Distribution of female deaths using the non-adjusted number and the adjusted number of
female deaths, using 2001 data. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
6.3 Distribution of female inhabitants by age classes in Arroios using 2011’s Census data,
and using the model without migrations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
6.4 Projected total of female inhabitants in Arroios using the model without migrations in 2016,
2021, 2026, 2031 and 2036.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
6.5 Projected distribution of female inhabitants by age classes in Arroios using the model
without migrations in 2016, 2021, 2026, 2031 and 2036. . . . . . . . . . . . . . . . . . . . 41
6.6 Distribution of female inhabitants by age classes in Arroios using 2011’s Census data,
and using the model with one migratory flow. . . . . . . . . . . . . . . . . . . . . . . . . . 42
6.7 Projected total of female inhabitants in Arroios using the model with one migratory flow in
2016, 2021, 2026, 2031 and 2036. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
6.8 Projected distribution of female inhabitants by age classes in Arroios using the model with
one migratory flow in 2016, 2021, 2026, 2031 and 2036. . . . . . . . . . . . . . . . . . . . 43
6.9 Projected distribution of female inhabitants by age classes in Arroios using the model with
one migratory flow in 2016, 2021, 2026, 2031 and 2036. . . . . . . . . . . . . . . . . . . . 46
6.10 Estimated internal and external flows of female inhabitants by age classes in Arroios using
the model with internal and external flows. . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
6.11 Distribution of female inhabitants by age classes in Arroios using 2011’s Census data,
and using the model with Leslie matrix that includes immigration and emigration I. . . . . 50
6.12 Projected total of female inhabitants in Arroios using the model with Leslie matrix that
includes immigration and emigration I in 2016, 2021, 2026, 2031 and 2036. . . . . . . . . 51
6.13 Projected distribution of female inhabitants by age classes in Arroios using the model with
Leslie matrix that includes immigration and emigration I in 2016, 2021, 2026, 2031 and
2036. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
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6.14 Distribution of female inhabitants by age classes in Arroios using 2011’s Census data,
and using the model with Leslie matrix that includes immigration and emigration II. . . . . 53
6.15 Projected total of female inhabitants in Arroios using the model with Leslie matrix that
includes immigration and emigration II in 2016, 2021, 2026, 2031 and 2036. . . . . . . . . 54
6.16 Projected distribution of female inhabitants by age classes in Arroios using the model with
Leslie matrix that includes immigration and emigration II in 2016, 2021, 2026, 2031 and
2036. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
6.17 Projected distribution of female inhabitants using models without migration and with one
migratory flow in 2016, and distribution of female inhabitants of the population sample
taken from 2017, by age classes in Arroios. . . . . . . . . . . . . . . . . . . . . . . . . . . 58
6.18 Projected distribution of female inhabitants using models with Leslie matrix that includes
immigration and emigration I and II in 2016, and distribution of female inhabitants of the
population sample taken from 2017, by age classes in Arroios. . . . . . . . . . . . . . . . 59
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Chapter 1
Introduction
1.1 Motivation
The knowledge of how a population will evolve over the years is essential to decision makers, since
they can create and follow strategies accordingly.
It can be achieved by forecasting a population, i.e. by making predictions based on current and past
data. Then, one can be aware of certain intrinsic aspects of an ecosystem, such as: the potential growth,
the age distribution of its individuals, survivors and descendants, fertility and mortality rates. Nonethe-
less, the projection of these intrinsic parameters is in fact conditional to one or more assumptions which
can indeed change. Sensitivity analysis measures how these aspects will react to perturbations (migra-
tion for example).
Assuming that age of individuals is known, there are two main approaches to exploit the dynamics
of the population in question: using a continuous-time integral equation, first introduced by F. R. Sharpe
and A. J. Lotka [1]; or developing a matrix formulation with age classes, presented by H. Bernardelli [2],
E.G. Lewis [3] and by P.H. Leslie [4].
In order to properly study population dynamics, models using Leslie matrices must be applied, based
on current and past data found at the Census, or other databases.
This thesis is closely related to a project called Conhecer Arroios, which was a partnership between
Junta de Freguesia de Arroios and Instituto Superior Tecnico de Lisboa. The main goal of this project
was to know how Arroios, a parish situated in Lisbon with more than 30000 inhabitants, was evolving
with time and predict the population growth in the future, by analysing 2001 and 2011 Census’ data and
by retrieving intercensus data from a population sample in 2017. In fact, I was coauthor of this study [5],
which conveyed the analysis of the evolution of Arroios parish along time and its projections, based on
2001 and 2011 Census’ data, and a population sample retrieved in 2017.
Thus, the decision makers can invest on infrastructures and services with guidance about the future
needs of their citizens. Otherwise, the lack of knowledge about the forecasted residents in a parish,
or even in a country, could lead to unnecessary expenditure and to depreciation of the domains on
demand at a later time. Therefore, in order to forecast useful pointers about the population’s future
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behaviour, useful and effective models must be found based on the current inhabitants. Also, there is
the need to analyse the response or sensitivity of evolutionary entropy to perturbations in the microscopic
parameters.
Another subject related to the predicted models is how to compress a Leslie matrix and, consequently
these models. That is how to, from a 5 year-age class matrix, achieve a 10 year-age class matrix without
any further input, by instance. Therefore, the applied model can be adapted according to the focus of
the study in question without needing to recalculate every parameter for every age-class, which can lead
to a reduction of the error produced by the iteration of the model.
1.2 Claim of Contributions
In what follows, we can point several contributions of our work.
• We propose several models that include migration flows;
• We analyse and applied several measures of evolutionary entropy-sensitivity to Arroios parish;
• We develop an algorithm that allows the compression of Leslie matrices;
• We implement the models and measures of evolutionary entropy in a real world context.
1.3 Thesis Outline
We conclude by presenting a brief overview of this dissertation.
In Chapter 2, the background, regarding the concepts and relevant theorems, and the state of the art
of population models applied before, are explored.
In Chapter 3, we develop and explain population models where the first two, model without migratory
flows and model with one migratory flow, can be already found in literature, and the remaining ones
contemplate the need for accounting both immigration and emigration.
In Chapter 4, we approach the partial derivatives of evolutionary entropy, deriving the characteristics
of the evolutionary entropy, and analysing the growth rate sensitivity.
In Chapter 5, the algorithm allowing the compression of the Leslie matrices is detailed.
In Chapter 6, we present and discuss the results given a real world context, obtained by applying the
models and measures, explained on previous Chapters, to a parish in Lisbon called Arroios.
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Chapter 2
Background
In this chapter we introduce some fundamental concepts and theorems related to matrix population.
Also, we introduce the Leslie and Lefkovitch matrices. A number of applied models by several authors
is presented.
2.1 Concepts and relevant theorems
2.1.1 Matrices
Definition 2.1. A square matrix, called A, is reducible iff it can be placed into block upper-triangular
form by simultaneous row/column permutations.
A square irreducible matrix is a non-reducible matrix. [6] That is, an irreducible matrix is a square
nonnegative matrix [7] such that
∀i,j∃k>0 : Ak(i, j) > 0. (2.1)
Definition 2.2. A square primitive matrix Aij [7] is a nonnegative matrix where some power of A = (i, j)
is positive.
Theorem 2.3. A positive square matrix is primitive and a primitive matrix is irreducible.[8]
Definition 2.4. A right eigenvector XR [9] is a column vector verifying
AXR = λRXR (2.2)
Notice that the term eigenvector is used to refer to a right eigenvector.
Definition 2.5. A left eigenvector XL [9] is a row vector verifying
XLA = λLXL (2.3)
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Theorem 2.6. (Perron-Frobenius Theorem) If all elements aij of an irreducible matrixA are nonnegative,
then
R = min{Mλ}. (2.4)
is an eigenvalue of A and all the eigenvalues of A lie on the disk
|z| ≤ R, (2.5)
where, if λ = (λ1, ..., λn) is a set of nonnegative numbers (which are not all zero),
Mλ = inf{µ : µλ >n∑j=1|aij |, 1 ≤ i ≤ n}. (2.6)
Furthermore, if A has exactly p eigenvalues (p ≤ n) on the circle |z| = R, then the set of all its
eigenvalues is invariant under rotations by 2πp about the origin. [9]
2.1.2 Graphs
A graph [10] is a collection of points, called vertices or nodes, that can be connected using lines,
called edges or arcs. There are three main types of graphs:
• simple graphs: there is at most one edge connecting two vertices;
• multigraphs: two vertices can have more than one edge connecting them, without any loops (self-
connected vertices);
• pseudographs: multigraphs that allow loops.
If a graph has labelled edges and vertices, it is called a labelled graph. Otherwise, it is an unlabelled
graph.
If the edges of a graph are undirected, then it is called an undirected graph. Otherwise it can be
a directed graph (the edges have arrows representing a unilateral or bilateral direction), or an oriented
graph (directed graph in which each edge has a unique direction).
Each edge can also have a weight associated. In that case, the graph is called weighted graph.
Otherwise, it is called unweighted graph.
A graph can be represented using an adjacency matrix, which is a matrix with rows and columns
labelled by graph vertices with 0 or a number different from 0 that represents the weight of an edge
connecting the vertices vi and vj . If the matrix has a 0 on position (vi, vj) then vi and vj are not
connected, that is do not have any edge between them. Otherwise, the value on (vi, vj) is the weight of
the edge connecting vi and vj .
A symmetric adjacency matrix represents an undirected graph and loops are represented with values
different from 0 on the matrix diagonal.
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2.1.3 Population matrices
Next we will exploit two types of population matrices: Leslie and Lefkovitch matrices. Due to the
regularity of the age intervals in the accessed data, there is no need to use the Lefkovitch matrix.
Instead, in this thesis we will use the Leslie matrix.
Leslie Matrix
Leslie Matrix rises as a possible answer for how are the survivors’ and descendants’ ages of a
certain population distributed, considering successive intervals of time and supposing that the rates
of fertility and mortality are applied to all elements equally and constant over time. The details are
further developed by H. Leslie [11] and H. Caswell [12]. In this model, only the female population will be
studied, being expressed by m + 1 linear equations organised in age classes. From now on, we adopt
the convention used on H. Leslie’s article [4].
Let:
• nxtbe the number of females alive in the age group x to x+ 1 at time t;
• Px be the survival probability of a female from an age class between x and x + 1 at time t to the
next age class x+ 1 to x+ 2 at time t+ 1. It has values between 0 < Px < 1;
• Fx ≥ 0 be the number of daughters born between t and t + 1 that are still alive in the age class
from 0 to 1 at time t+ 1 and whose mothers lived aged x to x+ 1.
Hence, the age distribution at the end of one unit interval can be expressed as:
m∑x=0
Fxnx0 = n01 (2.7)
P0n00 = n11 (2.8)
P1n10 = n21 (2.9)
P2n20 = n31 (2.10)
. . . (2.11)
Pm−1nm−10 = nm1 , (2.12)
which can be rewritten as the following m + 1 square matrix L × n0 = n1, where n0 and n1 are column
vectors providing the age distribution at t = 0, 1:
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L =
F0 F1 F2 . . . Fm−2 Fm−1 Fm
P0 0 0 . . . 0 0 0
0 P1 0 . . . 0 0 0
0 0 P2 . . . 0 0 0...
......
. . ....
......
0 0 0 . . . Pk 0 0
0 0 0 . . . 0 Pm−1 0
. (2.13)
If Fm = 0, the determinant of the matrix L is 0. Hence, the L is singular. Moreover, the age distribution
at time t is given by
Lt ×
n00
n10
n20
...
nm0
, (2.14)
that is, the multiplication between the age distribution at t = 0 and the matrix Lt. Then, the number of
female alive elements in the population at time t can be calculated as:
nj−10 ×m∑i=1
Lti,j . (2.15)
We are now going to derive the elements Fx and Px of the matrix.
Notice that, if there are nx,0 women alive in the age class from x to x + 1 at time t = 0, then the
survivors of this class will be the females in x+ 1 to x+ 2 age class at time t = 1. Then,
Pxnx,0 = nx+1,1. (2.16)
For Px’s derivation (Px is the survival probability of a female from an age class between x and x+ 1
at time t), we have:
Px = Lx+1
Lx, (2.17)
where Lx and Lx+1 is the number of females alive in the age group x to x + 1 and x + 1 to x + 2,
respectively, in the data collected from several databases.
Consider now Fx, which is the number of daughters born between t and t+ 1 that are still alive in the
age class from 0 to 1 at time t+ 1 and whose mothers lived aged x to x+ 1.
Let:
• mx be the fecundity per-capita for females of age class x;
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• lx = p0p1...px−1 be the survival from class 0 to x− 1.
Then we can calculate Fx:
Fx = Pxmx+1. (2.18)
Lefkovitch Matrix
A Lefkovitch matrix, explained and applied in several ecological studies on Lefkovitch’s article [13],
is a squared matrix describing populations with stage or size structure, where:
• Fx is the number of daughters that survived from age class 0− 1 and whose mothers are from age
class x to x+ 1;
• Px,x is the survival probability of females of the age class x in time t that still belong to class x in
time t+ 1;
• Px,x+1 is the survival probability of females from age class x in time t to next age class x + 1 in
time t+ 1
Thereby we obtain the following matrix:
L =
F0 + P0,0 F1 F2 . . . Fm−2 Fm−1 Fm
P0,1 P1,1 0 . . . 0 0 0
0 P1,2 P2,2 . . . 0 0 0
0 0 P2,3. . . 0 0 0
......
.... . .
......
...
0 0 0 . . . Pm−2,m−1 Pm−1,m−1 0
0 0 0 . . . 0 Pm−1,m Pm,m
. (2.19)
Notice that Px,x+Px,x+1 gives the total survival rate for the age class x and that, since newborns can-
not reproduce themselves, F0 + P0,0 = P0,0. Also, observe that the diagonal entries give the probability
of a female remaining in the same class from year t to t+ 1.
A pseudo-Leslie matrix is a matrix that can be decomposed in a row matrix (does not need to be a
non-negative and a subdiagonal positive matrix). Also, Lefkovitch matrices are similar to Leslie matrices
as pseudo-Leslie matrices. [14]
2.1.4 Relation between Leslie Matrices and Graphs
Notice that Lt in 2.13 can be seen as an adjacency matrix of a oriented and weighted graph. There-
fore, another way of representing a Leslie matrix Gc can be:
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C0 C1 C2 Cm−2 Cm−1 Cm
F0
P0 P1
F1F2
Pm−2
Fm−2
Pm−1
Fm−1
Fm
. . .
2.1.5 Population Dynamics
Evolutionary Entropy
Definition 2.7. Evolutionary entropy H is a statistical parameter that describes the diversity of pathways
of energy flow between the elements that compose the microlevel (e.g. individuals in the population),
and characterises the rate at which macroscopic variables, after a random perturbation, return to their
steady-state condition, that is, the robustness or stability of the hierarchy.
Note that this statistical measure is positively correlated with the elements that compose the mi-
crolevel.
Furthermore, evolutionary entropy can be defined as the rate at which the stochastic process P
generates information.
Analytically,
H =d∑i=1
πiHi (2.20)
with
Hi = −n∑j=1
pij log pij . (2.21)
Hi is called the Shannon-entropy associated with state X of the Markov Chain.[15]
Moreover, H is the weighted average, taken over all the stationary states πi, of Hi. Thus,
H =d∑i=1
πiHi = −n∑j=1
πipij log pij (2.22)
Robustness
Definition 2.8. The robustness of the system R is the fluctuation decay rate of Pε(t) on a logarithmic
time scale and characterises the insensitivity of a system to changes in the microlevel parameter.
Note that the entropy H and the robustness R are positively correlated.
Theorem 2.9. (Entropy-Robustness Theorem) Let ∆H = H ∗−H and ∆R = R∗−R be the change on
entropy and robustness, respectively, resulting from a perturbation in the parameters that describe the
network. Then
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∆H ×∆R > 0. (2.23)
Therefore, this theorem states that an increase in entropy leads to an increase in robustness. Hence,
the system is more insensitive to changes in the microlevel parameters that describe the network.[15]
Resilience
Definition 2.10. The resilience of the system is an aggregate property that depends on the response of
entropy to perturbations of the various linkages that compose the network.
Sensitivity
Definition 2.11. The sensitivity Sij relative to the changes of a system defined by a squared matrix Aij
is given by the derivatives of λ respective to the entries of A, where λ corresponds the eigenvalues of
A. That is,
Sij = ∂λ
∂aij= viuj , (2.24)
where vi and uj are the components of the left and right dominant eigenvectors, which are always
positive. [16]
Elasticity
Definition 2.12. The elasticity eij of a system defined by a squared matrix Aij [15] is given by
eij = aij∂r
∂aij= aij
λSij , (2.25)
where
r = log λ. (2.26)
2.2 State of the art
The idea of projecting a structured population size via a model with discrete time steps was intro-
duced by E.G. Lewis [3] and P.H. Leslie [4]. After that, some extensions of these basic models were
published by M.H. Williamson [17], M.B. Usher [18], L.P. Lefkovitch [13], C.A. Bosch [19], and recently
J.F. Alves and H. Oliveira [14].
In order to study populations in more detail, we need to decide upon the structure of the population
model. That is we need to choose the variables that confine all the previous data of an inhabitant that are
important to predict his future, as explained on Metz and Diekmann [20], and on Chapter 3 of H.Caswell’s
book [12].
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As H. Caswell pointed out in his article [21], it has been shown that it is beneficial to opt for age and
stage as the variables, since these models allow to take more information about the population than the
ones that rely only on stage as a variable. Also, if the vital rates are dependent on age and stage, then
it becomes essential to explore its correlation and therefore, the model must rely on these two variables.
Notice that stage is a relevant criteria to that population, for example, size, physiological condition,
fertility, or even spatial location, as seen on A.Rogers’ article [22].
Population growth, age and stage structure, and reproductive value belong to population dynamics,
while cohort dynamics refer to survivorship, life expectancy, age at death and generation time. Both pop-
ulation and cohort dynamics are studied calling upon a matrix model. As the name indicates, population
dynamics depends on society as a whole, including deaths and born individuals, while cohort dynamics
depends on individuals by themselves.
Since we are dealing with Human populations, we will classify individuals into the discrete stages
age and size. Population dynamics, projecting the population from time t to t + 1, can be showed as a
square matrix At as follows:
nt+1 = At × nt (2.27)
where nt+1 and nt are vectors with entries nit+1 and nit representing the number of individuals in a
certain stage i at time t+1 and t, where t+1 is the next iteration of t. The individuals can be classified into
age classes, done in Spain’s projection by H. Caswell and N. Sanchez Gassen [23], or the reproduction
for example.
We can classify these models into two categories based on the nature of At:
1. aij(t) are constants: the matrix population model is linear and time-invariant.
2. aij(t) are not constants: the matrix population model is non-linear and density-dependent.
In this thesis, only linear and time-invariant models are approached.
These models may have a change on the entries aij(t) due to external environmental periodic or
stochastic parameters, or to internal density or frequency dependence, as explained by H. Leslie on
[24].
Notice that a population modelled by a matrix with these characteristics evolves according to a expo-
nencial growth rate which is given by the maximum eigenvalue of At, where At is supposed irreducible
and primitive.
If the individuals are only subdivided into age classes then we use a Leslie matrix to model the
population. This approached was followed by P.H. Leslie [4].
Otherwise, if we want other criteria, such as size or stage, to classify the individuals, we must use a
Lefkovitch matrix.
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2.2.1 Model proposed by H. Leslie [11]
The growth rate of a population can be achieved by finding the eigenvalues and eigenvectors of the
matrix L, explained on 2.1.3, which correspond to population growth rate, stable life distribution and
reproductive value.
Let the population model be written as:
n(t+ 1) = L× n(t). (2.28)
Then, if we want to find the eigenvalues we solve:
(L− λ Id) n(t) = 0. (2.29)
Notice that the roots of the equation can be complex numbers.
The growth rate of a certain population λ is the eigenvalue that has the largest absolute value and it
will determine the expected behaviour of the population in projections. The other eigenvalues determine
the dynamics of the population. Also, the right eigenvector represents the stable age distribution, while
the left one describes the reproductive value.
2.2.2 Model proposed by L.P. Lefkovitch [13]
Let n(t) represent the column vector of a certain female population at time t subdivided by age
classes and the matrix L as presented on 2.1.3. Then
n(t+ 1) = L× n(t), (2.30)
that is, the distribution of the population at the next time step t + 1 is given by the product of the L and
n(t), the population at time t.
2.2.3 Model proposed by H. Caswell and N. Sanchez Gassen [23]
In [23] of H. Caswell and N. Sanchez Gassen, it was applied a model where the Leslie matrix used
has a particularity: it accounts for the emmigration when calculating the survival rate for each age-
class. That is, the survival rate is obtained by subtracting to 1 the mortality and the emigration to foreign
countries for each age-class. Also, a column vector was added to the model, representing immigrations
from foreign countries distributed by age-classes. Hence, the model can be written as:
n(t+ 1) = L× n(t) + Cimmig(t) (2.31)
with n(t = 0) = n0 and where n(t+ 1) is the vector of the female population subdivided by age-classes
at time t + 1, L is the Leslie Matrix with emigration calculated at time t, n(t) is the vector of the female
population subdivided by age-classes at time t + 1, and Cimmig(t) is the column vector containing the
immigrations divided by age-classes.
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This model does not adapt to the population inside a parish because there is the need to account not
only for flows coming from and to foreign countries, but those from and to the same country and different
parishes.
2.2.4 Population Dynamics
A crucial analysis of the population dynamics must be made in order to know how a perturbation can
affect the intrinsic variables of the population in study.
Thus, some analysis on the evolution of biological aging, i.e. the increase in mortality after maturation
of an organism, can be made. In fact, W.D. Hamilton, in [25], reported that higher fertility will lead to
increases in biological aging unless the resulting extra mortality happens only in immature ages. This
conclusions lead to the age-classified life cycles.
L. Demetrius showed in [26] that, for populations characterized by a stable size or by small fluc-
tuations of it, an unidirectional increase in population entropy for a large period of time leads to an
evolutionary change caused by mutation and natural selection. Also, for populations who experienced
an exponential growth, the same effect can be achieved by an unidirectional increase in growth rate and
a decrease in entropy for periods of time.
The competition between individuals for limited resources can be seen in two ways: classically, where
the process is deterministic and where its prediction relies only on the growth rate of the several popu-
lations (Malthusian selection principle); and non-classically, where the process is considered stochastic
and deeply related to the populations size and their evolutionary entropy and also by the availability of
the resource in question. As explored in L. Demetrius and S. Legendre [27], and in L. Demetrius [28],
the non-classical view which they call entropic selection principle, encapsulates two main contrasting
relations, where variants will have a selective advantage and an increase in frequency if:
• the populations have higher entropy and the resources are limited, constant and diversified;
• the populations have lower entropy and the resources are singular and suffer from disparities of
availability.
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Chapter 3
Proposed Models
This chapter is mainly focused on presenting the proposed models which convey closed systems
and systems with migration, we are designing models for real world application, namely to the parish of
Arroios in Lisbon where the field work was performed.
All models use a Leslie matrix instead of a Lefkovitch matrix since we are considering regular age
classes, as it is presented on Census data, and, by doing so, we simplify the model.
Moreover, given that only women can produce new human beings and their number is highly corre-
lated to the growth of a population, we shall use a single gender projection, namely the projection of the
female individuals, as employed in the literature [29].
3.1 Model without migratory flows
The model without migratory flows, considered on H. Leslie’s article [4], is written as:
n(t+ 1) = L× n(t), (3.1)
with n(t = 0) = n0 and where n(t+ 1) is the vector of the female population subdivided by age-classes
at time t + 1, L is the Leslie Matrix, in 2.1.3, calculated at time t and n(t) is the vector of the female
population subdivided by age-classes at time t+ 1.
This model is the simplest presented on this thesis. It can only be applied to closed systems, that is,
systems without any kind of flows, e.g. immigration and emigration.
Therefore, if we consider the Arroios parish, this model won’t be very well adjusted to its population,
since these ignored flows are very relevant and of high impact on the overall projection.
3.2 Model with one migratory flow
To cover the migration flows and based on 3.1, a Leslie matrix with the intrinsic population (consider-
ing a closed system) adds to a column vector with the proportions of the flows (subtraction of immigration
to emigration). That is:
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n(t+ 1) = L(
Id + α diag fluximm−em)
n(t), (3.2)
with n(t = 0) = n0 and where n(t+ 1) is the vector of the female population subdivided by age-classes
at time t + 1, L is the Leslie Matrix calculated at time t, n(t) is the vector of the female population
subdivided by age-classes at time t + 1, fluximm−em is the proportion of the flux relatively to the total
female residents and α is the adjustment factor of the flows along the years.
The main disadvantage of this model is that we have no separate control over the immigration and
emigration, since the only column vector contains both of them.
3.3 Model with immigration and correction flows
This model allows us to take control over the immigration and correction flows, solving in some way
the issue found with 3.2.
It can be written as:
n(t+ 1) = L(
Id + α diag fluximm + β diag fluxcor)
n(t), (3.3)
with n(t = 0) = n0 and where n(t+1) is the vector of the female population subdivided by age-classes at
time t+1, L is the Leslie Matrix calculated at time t, n(t) is the vector of the female population subdivided
by age-classes at time t+ 1, fluximm is the proportion of the immigration flux relatively to the projected
total female residents and α is its adjustment factor along the years, whilst fluxcor is the proportion
of a correction vector that includes emigration and errors in estimation of immigration relatively to the
projected total female residents and β is its adjustment factor along the years.
However, internal immigration, between parishes in the same country, is less prone to fluctuations
comparing to external immigration, between two different countries. The same applies for internal and
external emigration. Also, the vector fluximm was obtained based on the multiplication of the proportion
of Portugal’s female external immigrants for each age class, by the total number of female inhabitants in
Arroios that didn’t live in this parish five years ago. Consequently, some age classes may be predicted
having more immigration than the one observed in reality, being compensated by the opposite scenario
(higher emigration than in reality), since the total number of immigrants in Arroios is available to us.
Therefore, each of them has different rates and different adjustment factors along the years, even
though they are considered in the same column vectors and with the same adjustment factors in this
model, fluximm and fluxcor.
3.4 Model with internal and external flows
We can consider two types of migration: internal, between parishes in the same country, and exter-
nal, coming from or going to another country. These flows experience different fluctuations with time.
The external flows depend on various factors, for example the economy and working conditions of both
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countries. Nonetheless, the factors between parishes of the same country do not differ greatly through-
out the years and so, internal flows are more stable. In other words, external flows are more prone to
change when considering 10 years, by instance.
Hence, we can consider the following model:
n(t+ 1) = L(
Id + α diag fluxint + β diag fluxext)
n(t), (3.4)
with n(t = 0) = n0 and where n(t+ 1) is the vector of the female population subdivided by age-classes
at time t + 1, L is the Leslie Matrix calculated at time t, n(t) is the vector of the female population
subdivided by age-classes at time t+ 1, fluxint is the proportion of the internal flux relatively to the total
female residents and α is its adjustment factor along the years, whilst fluxext is the proportion of the
external flux relatively to the total female residents and β is its adjustment factor along the years.
This model allows us to adjust our projections according to the expected internal and external flows,
while giving us pointers to the impact of migrations to the intrinsic population in study. In this way, we
can also test what may happen to the inhabitants if there is an increase or decrease in migration.
3.5 Model with Leslie matrix that includes immigration and emi-
gration I
In order to study the sensitivity analysis of the inhabitants including the ones that are subject to the
migration flows, there is the need to place the column vectors mentioned on 3.4 inside the Leslie matrix.
That is, to account the internal and external flows in the survival probability of a female from one class
to the next one, and to account for the fertility rates of the total population, which includes the intrinsic
inhabitants and the ones coming from the migration flows.
On that account, we obtain the following Leslie matrix:
L′ =
F ′0 F ′1 F ′2 . . . F ′m−2 F ′m−1 F ′m
P ′0 0 0 . . . 0 0 0
0 P ′1 0 . . . 0 0 0
0 0 P ′2 . . . 0 0 0...
......
. . ....
......
0 0 0 . . . P ′k 0 0
0 0 0 . . . 0 P ′m−1 0
, (3.5)
where F ′x ≥ 0 is the number of daughters born between t and t + 1 that are still alive in the age class
from 0 to 1 at time t+ 1 and whose mothers lived aged x to x+ 1 and were an inhabitant or a immigrant
at time t + 1, and P ′x is the survival probability plus the migration flows of a female from an age class
between x and x+ 1 at time t to the next age class x+ 1 to x+ 2 at time t+ 1.
The model is:
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n(t+ 1) = L’× n(t). (3.6)
With this model, we can study the influence of changing the parameters in the population with migra-
tion and how much it alters from the population without flows.
Computationally, this method uses immigration and emigration flows resulting from the model with
internal and external flows, described on 3.4, and integrates them into the Leslie matrix by performing a
proportion that reduces to half the time span that these refer to. After, two time steps (that is 10 years),
the number of daughters resultant from this altered matrix is obtained and compared to the first entry
of the population. In this way, new proportions for the fertility rates containing all female inhabitants are
revealed.
3.6 Model with Leslie matrix that includes immigration and emi-
gration II
This model bears in mind 3.5 but follows a different computacional approach.
Therefore, the Leslie matrix is given by:
L′′ =
F ′′0 F ′′1 F ′′2 . . . F ′′m−2 F ′′m−1 F ′′m
P ′′0 0 0 . . . 0 0 0
0 P ′′1 0 . . . 0 0 0
0 0 P ′′2 . . . 0 0 0...
......
. . ....
......
0 0 0 . . . P ′′k 0 0
0 0 0 . . . 0 P ′′m−1 0
, (3.7)
where F ′′x ≥ 0 is the number of daughters born between t and t + 1 that are still alive in the age class
from 0 to 1 at time t+ 1 and whose mothers lived aged x to x+ 1 and were an inhabitant or a immigrant
at time t + 1, and P ′′x is the survival probability plus the migration flows of a female from an age class
between x and x+ 1 at time t to the next age class x+ 1 to x+ 2 at time t+ 1.
We obtain the following model:
n(t+ 1) = L”× n(t). (3.8)
This method can be computationally described as follows. First, we add unknowns parameters to
the survival rates Pi, from the Leslie matrix without any migration explained in 3.1. Then we obtain the
resulting vector of the population after two time steps (i.e. 10 years), and, as in 3.5, we compare the
number of daughters resultant from this altered matrix and the first entry of the population. Afterwards,
we put an unknown in the first entry of the matrix F ′′0 in order to be able to calculate a system with all the
unknowns. Since no daughters with less than 4 years old can produce offsprings, the unknown placed
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in F ′′0 is not correct when taken in practice. Therefore, the value obtained for this unknown is distributed
between all non-zero entries referring to fertility according to their previous proportion between each
other, and was placed a 0 in the first entry of the matrix.
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Chapter 4
Partial Derivates of Evolutionary
Entropy
Evolutionary entropy describes the diversity of pathways of energy flows between the elements that
compose the microlevel. That is, evolutionary entropy is a function of the change caused by perturba-
tions on a system that can be a biological, a metabolic or even an economic system. It allows for the
specification of robustness, the rate at the macroscopic variables that constitute the system return to
their steady state after a random perturbation.
The next sections will lead to the analysis of the sensitivity of evolutionary entropy to perturbations
in the microscopic parameters that describe the system.
This is based on L.A. Demetrius [15], and on H. M. Oliveira and L.A. Demetrius [30].
4.1 Evolutionary Entropy
Consider the following irreducible incidence matrix, associated with a graph,
A = (aij) ≥ 0. (4.1)
By the Perron-Frobenius theorem in 2.6, A has a dominant eigenvalue λ, and corresponding right
and left eigenvectors u = (u1, . . . , ud) and v = (v1, . . . , vd), respectively, such that:
Au = λu (4.2)
vA = λv (4.3)
(u,v) = 1. (4.4)
Also, this theorem ensures the differentiability of λ respective to the entries of A.
In many models, the incidence matrix A is considered as the matrix that specifies the steady state
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of the dynamic system, where the vector
u(t) = {ui(t)} (4.5)
represents its phase state. Therefore, the system will progress according to the equation
u(t+ 1) = Au(t). (4.6)
Now, consider the diagonal matrix U
U =
u0 0 . . . 0 0
0 u1 . . . 0 0...
......
. . ....
0 0 . . . ud−1 0
0 0 . . . 0 ud
, (4.7)
and the stochastic matrix P = (pij) = aijujλui
can be represented by
P = 1λ
U−1AU. (4.8)
Notice that P describes a Markov process with transition rate (pij)and stationary distribution Π =
(πi), where ΠP = Π and πi = viui.
Therefore, the evolutionary entropy H, weighted average of Hi, is the rate at which P generates
information, which analytically translates into
H =d∑i=1
πiHi = −d∑
i,j=1πipij log pij , (4.9)
where
Hi = −d∑j=1
pij log pij . (4.10)
As a consequence, Hi is the Shannon-entropy associated with the state Xi of the Markov chain.
4.1.1 Characteristics of the Evolutionary Entropy
4.1.1.1. Evolutionary entropy and generation time If S is the measure of the number of replicative
cycles in the network and T is the cycle time (the mean return time of the Markov process associated
with the matrix P), the evolutionary entropy H can be displayed as:
H = S
T. (4.11)
Let us show the characteristic above. From now on, fix the vertex a ∈ A where X = (1, . . . , d). Also,
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let X be the set of all states such that
X = { [a, β1, . . . , βn−1, a] : a→ β1 → · · · → βn−1 → a where βi 6= a for all i, and n ≥ 1 } . (4.12)
A state a ∈ X is a path of the graph G, which starts and ends at a, and does not visit a inbetween.
The probability of a given cycle which starts at a is given by
pa = pβ1pβ1β2 . . . pβn−1βn. (4.13)
The number of replicative cycles associated with the network will then given by
S = −∑a∈X
pa
log pa. (4.14)
In the graphs described by a unique replicative cycle S = 0, as illustrated in the graph below.
1 2 d. . .
Figure 4.1: S = 0, where S is a measure of the number of replicative cycles.
S increases proportionally to the number of replicative cycles increases as represented below.
1 2 d. . .
Figure 4.2: S > 0, where S is a measure of the number of replicative cycles.
Let
T =∑a∈X
|a|pa. (4.15)
In this case, we have |a| = n, the length of the cycle, for n such that
a = [a, β1, . . . , βn−1, a]. (4.16)
The quantitiesH, measure of the rate at which the process is generating information, and S, measure
of the uncertainty in the length of a randomly chosen cycle, differ only in terms of the value T , as
demonstrated on L.A. Demetrius, V.M. Gundlach and M. Ziehe’s article [31]
H = S
T. (4.17)
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4.1.1.2. Variational principles and evolutionary entropy The matrix P = (pij) when is derived from
the interaction matrix A = (aij) ≥ 0 represents a fundamental feature of its network. If r = log λ and Φ
is the reproductive potential, we obtain:
r = Φ + S
T. (4.18)
Consider the set MA of all stochastic matrices P = (pij) which satisfy the property
aij = 0⇔ pij = 0. (4.19)
The parameter r = log λ satisfies a variational principle, as demonstrated by L. Arnold, L.A. Demetrius
and V. M. Gundlach in [32]. Then:
r = log λ = supP∈MA
∑i,j
πipij(log pij − log aij). (4.20)
Furthermore, the supremum in 4.20 can be obtained by the unique stochastic matrix P = (pij),
defined by the relation
pij = aijujλui
. (4.21)
Using the two equations above, we have:
r = log λ = H + Φ, (4.22)
where H is the evolutionary entropy and Φ is the reproductive potential given by
Φ =∑i,j
πipij log aij . (4.23)
Using 4.18, we can get the last equation as
r = Φ + S
T. (4.24)
4.1.1.3. Evolutionary entropy and robustness An increase in entropy entails an increase in robust-
ness and, therefore, a greater insensitivity of an observable to perturbations in the microlevel parameters
that describe the network.
Let Pε(t) denote the probability that the sample mean deviates from its unperturbed value by more
than ε at time t.
As L.A. Demetrius, V.M. Gundlach and G. Ochs showed in [33], as t increases, Pε(t) converges to
zero.
Robustness can be quantified by analysing deviations of the observables of the system following an
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instantaneous perturbation of the microlevel parameters.
More formally, robustness R characterizes the insensitivity of an observable to structural changes
in the microlevel parameter and it is defined as the fluctuation decay rate of Pε(t) on a logarithmic time
scale. It can be written as:
R = limt→∞
− logPε(t)t
. (4.25)
The entropy H and the robustness R are positively correlated, as was also shown in [33].
In this case, using the Entropy-Robustness theorem in 2.9, we have the values
∆H = H∗ −H and ∆R = R∗ −R (4.26)
which are the changes in the variables that provoke a change in the parameters that describe the net-
work.
By this theorem, an increase in entropy leads to an increase in robustness, and, consequently, to a
larger insensitivity to perturbations in the microlevel parameters that characterize the network behaviour.
4.2 Sensitivity Analysis
For example, if we consider demographic networks, then the growth rate experiences stronger effect
due to variations in the fertility rates in lower age-classes than in higher age-classes.
Recall equation 4.22:
r = log λ = H + Φ,
which can be written as
H = r − Φ. (4.27)
Next we will exploit the sensitivity of the growth rate r = log λ.
4.2.0.1. Growth rate sensitivity[30] The sensitivities Sij are given by
Sij = ∂λ
∂aij= viuj , (4.28)
where vi > 0 and uj > 0 are the components of the left and right dominant eigenvectors, respectively.
Consider the matrix A, complying with the conditions mentioned on 4.1 , and recall the concept of
entropy in 2.11: there are exactly d2 such derivatives
Sij = ∂λ
∂aij.
Given the differentials in Au = λu such that
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dAu +Adu = dλu + λdu. (4.29)
Recall that vu = 1, where v is the left eigenvalue. Using the matrix notation for the usual Euclidean
inner product, we obtain
vdAu + vAdu = vdλu + vλdu↔ vdAu + λvdu = vdλu + λvdu (4.30)
that is,
vdAu = vdλu. (4.31)
Then, the intended result follows:
Sij = ∂λ
∂aij= viuj . (4.32)
We construct the sensitivity matrix S as:
S = (viuj)d×d = v⊗ u =( ∂λ
∂aij
)d×d
. (4.33)
Using r = log λ, we get:
∂r
∂aij= 1λviuj = 1
λSij , (4.34)
and the concept of elasticity eij
eij = aij∂r
∂aij= aij
λSij . (4.35)
One interesting remark is that the sensitivity of the elasticity is closely related to the sensitivity of the
growth rate.
The sensitivity of the elasticity measures the effective perturbation on the growth rate weighted by
the size of each structure matrix entry.
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Chapter 5
Compression of the Leslie matrices
For the sake of simplification, we shall use Leslie graphs which have Lt as incidence matrix.
Imagine that we want to compress the graph represented on 2.2.3, with c as the interval of each
age-class considered into a graph with n×c as the interval of each age-class. Without loss of generality,
let’s use n = 2.
Therefore, we want to discover F ′0, F ′1, . . . , F ′m2 and P ′0, P ′1, . . . , F ′m2 , illustrated in the graph G2c below,
such that they correspond to the values directly obtained using the fertility and survival rates for a Leslie
matrix with 2c-age-classes.
C ′0 C ′1 C ′2 C ′m2 −2 C ′m
2 −1 C ′m2
F ′0
P ′0 P ′1
F ′1F ′2
P ′m2 −2
F ′m2 −2
P ′m2 −1
F ′m2 −1
F ′m2
. . .
Suppose that m is an even number. Notice that, to obtain F ′0, F ′1, . . . , F ′m2 and P ′0, P ′1, . . . , F ′m2 from a
Leslie matrix with c-age-classes, these weights must be written in order of the number of females alive in
each age group (nx0 , nx1 , . . . , nxm ), and in order of the coefficients of a Leslie matrix with c-age-classes
(F0, F1, . . . , Fm2
and P0, P1, . . . , Fm2
).
A fertility rate Fi can be obtained by:
Fi = #(daughters given by women in age-class i)#(female population in age-class i) , (5.1)
which means that, if we double the time-interval and also double the age-class c, we can obtain the
fertility rates for a Leslie matrix subdivided into 2c age-classes depicted above in the graph by:
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F ′j = #(daughters given by women in age-class 2j) + #(daughters given by women in age-class 2j+1)#(female population in age-class 2j) + #(female population in age-class 2j+1)
if 0 ≤ j ≤ m
2 − 1 (5.2)
and
F ′m2
= #(daughters given by women in age-class m)#(female population in age-class m) . (5.3)
Therefore, relating to the graph representing the Leslie matrix subdivided into c age-classes:
F ′j = F2j + F2j+1
nx2j + nx2j+1
if 0 ≤ j ≤ m
2 − 1
F ′m2
= Fm
(5.4)
(5.5)
.
The same way of thinking can be applied to achieve the values of the survival rates (considering that
the system is closed for migrations), that are calculated as follows:
Pi = 1− #(women that passed away in age-class i)#(midyear female population in age-class i) . (5.6)
Consequently,
P ′j = P2j + P2j+1
nx2j + nx2j+1
if 0 ≤ j ≤ m
2 − 1 (5.7)
If m was an odd number, then we would obtain, from the c age-class Leslie graph Gc, the following
graph for the 2c age-class Leslie matrix G′2c:
C ′0 C ′1 C ′2 C ′m+12 −3 C ′m+1
2 −2 C ′m+12 −1
F ′0
P ′0 P ′1
F ′1F ′2
P ′m+12 −3
F ′m+12 −3
P ′m+12 −2
F ′m+12 −2
F ′m+12 −1
. . .
and the weights representing the fertility rates in the graph represented above are given as, for 0 ≤ j ≤m+1
2 − 1:
F ′j = F2j + F2j+1
nx2j+ nx2j+1
. (5.8)
Mutatis mutantis, the survival rates, for 0 ≤ j ≤ m+12 − 2, are calculated as:
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P ′j = P2j + P2j+1
nx2j + nx2j+1
. (5.9)
5.1 Re-weighting the edges of the graph G2c
In order to perform the proposed compression of the Leslie graph Gc, we must find an algorithm
containing a function that allows the re-weighting of the edges with the least possible error. This function
must only depend on the coefficients presented on the matrix L. From a graph’s point of view, this
procedure can only depend on the current weight of the edge wi of Gc, and the number of female
individuals on the population subdivided into c age-classes.
The idea is to perform a mean of the two edges, i and i + 1, that are going to be joined with a
correction factor. To find this factor, g, and as consequence the weight of the corresponding edge w′j of
G2c, the following system must be solved in order of wi, wi+1, nxiand nxi+1 (notice that the weights of
the edges considered are always rates and ai is the numerator of these):
wi = ainxi
wi+1 = ai+1
nxi+1
.
w′j = wi + wi+1
g
(5.10)
(5.11)
(5.12)
By solving the system above, we achieved the following result for g:
g =(nxi
+ nxi+1)(wi + wi+1)nxiwi + nxi+1wi+1
. (5.13)
That is, the weight of the edge w′j of G2c, if m is even, is given by:
w′j = w2j + w2j+1
(nx2j+ nx2j+1)(w2j + w2j+1)
nx2jw2j + nx2j+1w2j+1
if 0 ≤ j ≤ m
2 − 1.
w′m2
= wm if such wm exists
(5.14)
(5.15)
If m is odd, the new weight is given by, for 0 ≤ j ≤ m+12 − 1:
w′j = w2j + w2j+1(nx2j + nx2j+1)(w2j + w2j+1)
nx2jw2j + nx2j+1w2j+1
if such wm exists. (5.16)
Notice that, even if the studied Leslie matrix included migrations, which wasn’t a closed system, the
same procedure could be applied to re-weight each edge. This happens because the migrations (migx)
are added to the sub-diagonal as follows
Pj = 1−mortj −migj , (5.17)
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that is the probability of survival of some class j to the next class j + 1 is obtained by subtracting to 1
the mortality rate (mortx) and the migration rate, where the migration rate is the emigration rate (emx)
minus the immigration rate (immx):
migx = emj − immj . (5.18)
In more detail, the probability of survival of some class j to the next class j + 1, in an open system,
is not only dependent on the mortality rate, but also on the migration flow (the proportion of inhabitants
leaving the system and the proportion of individuals deciding to be a part of the system). But this
information can be compressed the same way as P ′j in a closed system 5.9:
P ′j = P2j + P2j+1
nx2j + nx2j+1
.
Therefore, we can apply 5.14 and 5.16 in open systems as well.
5.2 Proposed Algorithm
Suppose that the input Leslie matrix, lc, has dimensions n × n (a Leslie matrix is always a square
matrix). The complexity with regard to its input of the inner While of this algorithm is given by O(dn2 e),
while the outer While has complexity O(2). Therefore, the complexity of this algorithm is O(2dn2 e).
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Algorithm 1 Compression of the Leslie Matrix
1: procedure LESLIECOMPRESSED(lc,p)
Input:lc, Leslie matrix subdivided into c age-classes in the considered initial year t, and p, thecorresponding column vector containing the number of females
Output:l2c, Leslie matrix subdivided into 2c age-classes
2: Solve[
(a2j−1+a2j)(nx2j−1 +nx2j
) == (w2j−1+w2j)g , w2j−1 == a2j−1
nx2j−1, w2j == a2j
nx2j, a2j−1, a2j , nx2j−1 , nx2j
, g]
3: g[nx2j−1 , nx2j, w2j−1, w2j ] = (nx2j−1 +nx2j
)(w2j−1+w2j)(nx2j−1w2j−1)+(nx2j
w2j) ;4: q = 1;5: r = {}6: l = {lc1,Table[lcj,j−1, j, 2,Dimensions[lc]1]7: while q ≤ Dimensions[l]1 do8: i = 19: m1 = {}
10: w = lq11: while i ≤ dDimensions[w]1
2 e do12: if i 6= dDimensions[w]1
2 e then13: if w2i−1 6= 0&&w2i 6= 0 then14: m1 = Append
[m1, w2i−1+w2i
g[p2i−1,1,p2i,1,w2i−1,w2i]
]15: else16: if w2i−1 == 0&&w2i 6= 0 then17: m1 = Append
[m1, w2i
]18: else19: if w2i−1 6= 0&&w2i == 0 then20: m1 = Append
[m1, w2i−1
]21: else22: m1 = Append
[m1, 0
]23: end if24: end if25: end if26: else27: if isEven
[Dimensions[w]1
]then
28: m1 = Append[m1, w2i−1+w2i
g[p2i−1,1,p2i,1,w2i−1,w2i]
]29: else30: m1 = Append
[m1, w2i−1
]31: end if32: end if33: i+ +34: end while35: r = Append[r,m1]36: q + +37: end while38: if isEven[Dimensions[w]1] then39: lc = Table[PadLeft[PadRight[{r2,u+1},
Ceiling[Dimensions[lc]]12 − u], Ceiling[Dimensions[lc]]1
2 ], {u, 0, Dimensions[r2]1 − 1}]
40: else41: lc = Table[PadLeft[PadRight[{r2,u+1},
Ceiling[Dimensions[lc]]12 − u], Ceiling[Dimensions[lc]]1
2 ], {u, 0, Dimensions[r2]1 − 2}]
42: end if43: lc = Prepend[lc, r1]44: end procedure
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Theorem 5.1. The proposed algorithm for the compression of Leslie matrices has null error.
Proof:
Consider the following two tables concerning the data that we are going to use.
Age-classes C0 C1 C2 . . . Cm
Number of children born whose mothers belong to this class a0 a1 a2 . . . am
Number of women that survived from age-class Ci to Ci+1 s0 s1 s2 . . . sm
Number of women of this class n0 n1 n2 . . . nm
Table 5.1: Table with the number of children born, of women that survived from age-class Ci to Ci+1,and of women by c age-classes.
Age-classes C ′0 . . . C ′m2
Number of children born whose mothers belong to this class a′0 = a0 + a1 . . . a′m2
= am
Number of women that survived from age-class Ci to Ci+1 s′0 = s0 + s1 . . . s′m2
= sm
Number of women of this class n′0 = n0 + n1 . . . n′m2
= nm
Table 5.2: Table with the number of children born, of women that survived from age-class Ci to Ci+1,and of women by 2c age-classes, related with the women subdivided by c age-classes.
Therefore, the Leslie matrix subdivided into c age-class is given by:
Lc =
f0 = a0n0
f1 = a1n1
. . . fm−1 = am−1nm−1
fm = am
nm
p0 = s0n0
0. . . 0 0
0 p1 = s1n1
. . . 0 0
0 0. . . 0 0
0 0. . . pm−1 = sm−1
nm−10
. (5.19)
We have two cases to consider:
a. m is even
Let us suppose that m is even.
The Leslie matrix subdivided into 2c age-classes, using the c age-classes’ data:
L2c =
f ′0 = a0+a1n0+n1
f ′1 = a2+a3n2+n3
. . . f ′m/2−1 = am−2+am−1nm−2+nm−1
f ′m/2 = am
nm
p′0 = s0+s1n0+n1
0. . . 0 0
0 p′1 = s2+s3n2+n3
. . . 0 0
0 0. . . 0 0
0 0. . . p′m/2−1 = sm−2+sm−1
nm−2+nm−10
. (5.20)
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Using as input Lc and nc =
n0
n1
n2...
nm
for the proposed algorithm, we obtain for the fertilities:
f ′′0 = (f0n0 + f1n1)(f0 + f1)(n0 + n1)(f0 + f1) =
( a0n0n0 + a1
n1n1)( a0
n0+ a1
n1)
(n0 + n1)( a0n0
+ a1n1
) = a0 + a1
n0 + n1= f ′0 (5.21)
f ′′1 = (f2n2 + f3n3)(f2 + f3)(n2 + n3)(f2 + f3) =
( a2n3n2 + a3
n3n3)( a2
n2+ a3
n3)
(n2 + n3)( a2n2
+ a3n3
) = a2 + a3
n2 + n3= f ′1 (5.22)
. . .
f ′′m2
= fm = f ′m2, (5.23)
and for the survival probabilities:
p′′0 = (p0n0 + p1n1)(p0 + p1)(n0 + n1)(p0 + p1) =
( s0n0n0 + s1
n1n1)( s0
n0+ s1
n1)
(n0 + n1)( s0n0
+ s1n1
) = s0 + s1
n0 + n1= p′0 (5.24)
p′′1 = (p1n1 + p2n2)(p1 + p2)(n1 + n2)(p1 + p2) =
( s1n1n1 + s2
n2n2)( s1
n1+ s2
n2)
(n1 + n2)( s1n1
+ s2n2
) = s1 + s2
n1 + n2= p′1 (5.25)
. . .
p′′m2 −1 =
(pm2 −1nm2 −1 + pm
2nm
2)(pm
2 −1 + pm2
)
(nm2 −1 + nm
2)(pm
2 −1 + pm2
) =
=(sm
2 −1
nm2nm
2 −1 +sm
2nm
2nm
2)(sm
2 −1
nm2 −1
+sm
2nm
2)
(nm2 −1 + nm
2)(sm
2 −1
nm2
+sm
2 −1
nm2
)=
=sm
2 −1 + sm2
nm2 −1 + nm
2
=
= p′m2 −1
(5.26)
which corresponds to the Leslie matrix L2c, as expected.
b. m is odd
Let m is odd and consider the following Leslie matrix subdivided into 2c age-classes, using the c
age-classes’ data.
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L2c =
f ′0 = a0+a1n0+n1
f ′1 = a2+a3n2+n3
. . . f ′(m + 1)/2−2 = am−3+am−2nm−3+nm−2
f ′(m + 1)/2−1 = am−2+am−1nm−2+nm−1
p′0 = s0+s1n0+n1
0. . . 0 0
0 p′1 = s2+s3n2+n3
. . . 0 0
0 0. . . 0 0
0 0. . . p′(m + 1)/2−2 = sm−3+sm−2
nm−3+nm−20
. (5.27)
Following the procedure using as input Lc and nc =
n0
n1
n2
. . .
n4
, we obtain for the fertilities:
f ′′0 = (f0n0 + f1n1)(f0 + f1)(n0 + n1)(f0 + f1) =
( a0n0n0 + a1
n1n1)( a0
n0+ a1
n1)
(n0 + n1)( a0n0
+ a1n1
) = a0 + a1
n0 + n1= f ′0 (5.28)
f ′′1 = (f2n2 + f3n3)(f2 + f3)(n2 + n3)(f2 + f3) =
( a2n3n2 + a3
n3n3)( a2
n2+ a3
n3)
(n2 + n3)( a2n2
+ a3n3
) = a2 + a3
n2 + n3= f ′1 (5.29)
. . .
f ′′(m + 1)/2−2 =(f(m + 1)/2−1n(m + 1)/2−1 + f(m + 1)/2n(m + 1)/2)(f(m + 1)/2−1 + f(m + 1)/2)
(n(m + 1)/2−1 + n(m + 1)/2)(f(m + 1)/2−1 + f(m + 1)/2) =
=(a(m + 1)/2−1n(m + 1)/2
n(m + 1)/2−1 + a(m + 1)/2
n(m + 1)/2n(m + 1)/2)( a(m + 1)/2−1
n(m + 1)/2−1+ a(m + 1)/2
n(m + 1)/2)
(n(m + 1)/2−1 + n(m + 1)/2)( a(m + 1)/2−1n(m + 1)/2−1
+ a(m + 1)/2
n(m + 1)/2)
=
=a(m + 1)/2−1 + a(m + 1)/2
n(m + 1)/2−1 + n(m + 1)/2
= f ′(m + 1)/2−2
(5.30)
and for the survival probability:
p′′0 = (p0n0 + p1n1)(p0 + p1)(n0 + n1)(p0 + p1) =
( s0n0n0 + s1
n1n1)( s0
n0+ s1
n1)
(n0 + n1)( s0n0
+ s1n1
) = s0 + s1
n0 + n1= p′0 (5.31)
. . .
p′′(m + 1)/2−2 =(p(m + 1)/2−1n(m + 1)/2−1 + p(m + 1)/2n(m + 1)/2)(p(m + 1)/2−1 + p(m + 1)/2)
(n(m + 1)/2−1 + n(m + 1)/2)(p(m + 1)/2−1 + p(m + 1)/2) =
=( s(m + 1)/2−1n(m + 1)/2−1
n(m + 1)/2−1 + s(m + 1)/2
n(m + 1)/2n(m + 1)/2)( s(m + 1)/2−1
n(m + 1)/2−1+ s(m + 1)/2
n(m + 1)/2)
(n(m + 1)/2−1 + n(m + 1)/2)( s(m + 1)/2
n(m + 1)/2+ s(m + 1)/2−1
n(m + 1)/2−1)
=
=s(m + 1)/2−1 + s(m + 1)/2
n(m + 1)/2−1 + n(m + 1)/2
=
= p′(m + 1)/2−2
(5.32)
which corresponds to the Leslie matrix L2c, as expected.
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Chapter 6
Real Word Context
This thesis was developed alongside with a project called Conhecer Arroios, where an intercensus
study, referring to the year 2017, of the population of Arroios was made. I was coauthor of this study [5],
which conveyed the analysis of the evolution of Arroios parish along time and its projections, based on
2001 and 2011 Census’ data, and a population sample retrieved in 2017. This was a great advantage
since it allowed us to apply the models presented on 3 to a real world context.
In the next sections, we will explain how we obtained the data to apply to our projections. After,
we will approach the results obtained by projecting each model on 3, exploring age distributions and, if
applicable, migration flows. Next, the effect of changes to the parameters that constitute the population
will be studied, using the obtained three Leslie matrices. Finally, a comparison between the data from a
population sample in 2017 and the real world application of our models will be made.
6.1 Construction of Arroios’ Leslie Matrices based on 2001 and
2011 data
In order to make several projections of the population in Arroios, we must obtain some possible
Leslie matrices based on 2001 and 2011 data. Our models take into account some indices derived from
Census 2001 and 2011, for instance fertility and mortality indices, immigration and emigration.
The data from the Census is very incomplete regarding the considered parish and the required
variables to build this model (e.g. emigration and immigration). Therefore, we made several estimations
which are detailed on the following subsections.
6.1.1 Fertility Indices
There are multiple indices to calculate fertility and we explore some beginning with less precision
and then go forward and reach to one that has enough precision for our study.
In our case, we are only interested on newborn females, since they are the ones renovating the
population.
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We can obtain the Gross Reproduction Rate (GRR):
GRRi =∑j
#(daughters given by women in age-class j in year i)#(midyear female population in age-class j in year i) , (6.1)
which gives the number of girls born per woman. However, if we take into account mortality among
female births, then we calculate the Net Reproduction Rate or Net Replacement Rate:
NRRi =∑j
#(daughters alive given by women in age-class j in year i)#(midyear female population in age-class j in year i) (6.2)
Since the estimated number of stillborn children in Arroios during the years of 2001 and 2011 is almost
0 (in 2001 there was one stillborn daughter), we shall procede the calculations of fertility based on the
Gross Reproduction Rate.
Let us suppose that the age-specific fertility is different from 0 between 15 and 49 years old.
Since there was no available data for the number of daughters per their mothers’ age-class, for
Arroios, our calculations were based under the assumption that the distribution of the proportion of
daughters and mothers of this area is similar to Lisbon’s distribution. After obtaining this value, we
applied a correction to the extrapolation because the estimate of total number of daughters born was
not equal to the number of female babies born for parish considered. That is, we multiplied the number
of daughters per their mothers’ age-class by a factor
α = #(total estimated number of daughters born)#(total number of daughters born)
. (6.3)
The following plot gives a more clear idea of the distribution shift in order to match the known number
of daughters born.
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Figure 6.1: Distribution of female newborns using the non-adjusted number and the adjusted number offemale newborns, using 2001 data.
6.1.2 Mortality
The procedure applied for finding the mortality rates for each age class was similar to the fertility
indices.
Let us consider the Gross Mortality Rate (GMR):
GMRi =∑j
#(women that passed away in age-class j in year i)#(midyear female population in age-class j in year i) . (6.4)
Therefore, GMRi gives us the proportion of women in age-class j that died comparing to the total
number of women in that age-class, considering the same year i.
However, since there is no available data to cover the number of deaths per age-class, we assumed
that the distribution of the proportion of mortalities is similar to Lisbon’s distribution, and then we applied
a correction of this value since the estimate of the total number of deaths was not equal to the number
of deaths for each parish. The correction applied may be expressed as:
α = #(total estimated of women that passed away)#(total of women that passed away)
. (6.5)
The corresponding shift in mortality is illustrated in the figure below.
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Figure 6.2: Distribution of female deaths using the non-adjusted number and the adjusted number offemale deaths, using 2001 data.
6.1.3 Survival Indices
Remember the concept of Leslie matrix given on 2.1.3:
L =
F0 F1 F2 . . . Fm−2 Fm−1 Fm
P0 0 0 . . . 0 0 0
0 P1 0 . . . 0 0 0
0 0 P2 . . . 0 0 0...
......
. . ....
......
0 0 0 . . . Pk 0 0
0 0 0 . . . 0 Pm−1 0
. (6.6)
is a Leslie matrix where Pi,i+1 is the survival index and Fi is the fertility rate for each age-class.
When we consider a closed system (without any migrations), the survival index Pi of this matrix was
calculated by subtracting to 1 the mortality rate for each age-class.
Otherwise, when the migration flow was considered inside the matrix, the survival index Pi was
obtained by subtracting mortality and emigration rates and adding the immigration rate to 1.
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6.1.4 Immigration
We first obtained the total female population that didn’t live in each parish 5 years ago, which was
accomplished by multiplying, for 2001 and 2011, the total female population by the given proportion of
female residents that didn’t live in each parish. Observe that this subpopulation contains portuguese
and foreign nationalities.
In order to separate the vectors containing immigration from foreign nationalities and immigration
from portuguese women, we multiplied the proportion of foreign women living in each parish by the total
female population that wasn’t a resident of that parish five years ago. It had to be assumed that the
immigration was uniformly proportional over time to the number of foreign women.
We fitted our data using a quadratic function because there is no data available for 2006, needed
for the construction of the model. Applying this expression, a value for the number of foreign immigrant
women was obtained.
Then, we subtracted the total female immigrant population by the previous value and, therefore, the
number of portuguese immigrants of this parish was calculated.
Due to the lack of data for the distribution of immigration by age-classes in the parishes, we inferred
using Portugal’s immigration data, which only considers women arriving from foreign countries. There-
fore, it is assumed that the internal immigration (women coming from other parishes in Portugal) follows
a similar distribution.
Consequently, a vector containing the immigration of women of foreign nationalities divided by age-
classes between 2001 and 2011, and a vector containing the immigration of portuguese women divided
by age-classes between 2001 and 2011, were calculated.
Afterwards, the immigration ratio was calculated based on the number of female inhabitants in 2001.
6.1.5 Emigration to Foreign Countries
Using a scale parameter which accounts for the population of each parish and Portugal, we multiplied
it by the Portuguese emigration of each year, assuming that the parish follows the distribution of Portugal
over the years. A similar assumption was made regarding the age-class distribution of emigration, where
the proportion of the cumulative distribution between 2008 and 2015 by age-class was used in order to
obtain the vector containing the emigration of women of foreign nationalities divided by age-classes,
between 2001 and 2011.
6.1.6 Error Estimate
We used a forecast error in order to evaluate the error of the predicted model, which is the difference
between the observed value and its forecast. That is:
error =
∑21i=1|nesti2011
− ni2011 |ni2011
21 (6.7)
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where nesti2001, and ni2011 , is the estimated, and the observed, distribution of the female population
of the age-classes i, respectively.
6.2 Application of the models
In 3, several models were presented, conveying closed systems and systems with migration. It is
important to establish a connection between the theoretical point of view and its application to the real
world, in order to evaluate and compare models and attain better projections.
Having as goal to decrease the error, and since the Leslie matrix calculated for 2001 is considered
constant over time, we consider all iterations beginning from 2011, that is:
n2011+5i = L2011i × n2011. (6.8)
Also, notice that we suppose, in our calculations, the emigration and immigration flows follow an
uniform distribution with time.
Moreover, since our models only consider women, we can obtain the total population by multiplying
the number of female inhabitants by 1.86, which is the observed proportion between women and the
total population in 2011.
6.2.1 Model without migratory flows
Following the model without migratory flows explained on 3.1, we present the values obtained for
each parameter of these matrices, considering a Leslie matrix for closed systems. Notice that only
age-classes of 5 and 10 years will be considered in this research.
Therefore, the parameters for Leslie Matrices are displayed in the following tables.
F3 F4 F5 F6 F7 F8 F9
Leslie Matrix I 0.0101563 0.0286868 0.0443657 0.0401565 0.0126653 0.00325448 0.0000779663
Table 6.1: Fertility rates obtained from 2001’s data without considering migratory flows.
P0 P1 P2 P3 P4 P5 P6
Leslie Matrix I 0.998069 0.999676 0.999902 0.999466 0.999733 0.999054 0.998966
P7 P8 P9 P10 P11 P12 P13
Leslie Matrix I 0.998043 0.997554 0.995478 0.99625 0.995094 0.991306 0.989036
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P14 P15 P16 P17 P18 P19
Leslie Matrix I 0.978306 0.960356 0.921155 0.859404 0.744179 0.573544
Table 6.2: Survival rates obtained from 2001’s data without considering migratory flows.
If we compare the distribution of female residents in Arroios found in 2011, using Census data, and
using this model without migrations, we can closely analyse the effect of migrations.
• The model without migrations underestimates the number of newborns and relatively young people
(until the age class of 45-49 years old), which can induce to the conclusion that the effects of
immigration are stronger than the ones felt from emigration in these age classes, where young
couples bring their children to live in Arroios;
• The model without migrations gives a close number to the reality found in 2011 between age
classes of 50 and 59 years old;
• The model without migrations overestimates the number of relatively old inhabitants (after 60 years
old), from which we can infer a stronger role of emigration comparing to immigration. This can be
due to lack of nursing homes in Arroios and exodus of the elderly to caregivers’ house.
The shift between the estimated female population using this model in 2011 and the actual female
population encountered in Census 2011 is pictured below, and the error estimate between these two
populations is 94.068%.
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Figure 6.3: Distribution of female inhabitants by age classes in Arroios using 2011’s Census data, andusing the model without migrations.
Next, we present projections, using this model for closed systems, obtained for 2016, 2021, 2026,
2031 and 2036.
Figure 6.4: Projected total of female inhabitants in Arroios using the model without migrations in 2016,2021, 2026, 2031 and 2036..
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The figure above shows that this model without migrations predicts a significant decrease in female
population from 2001 to 2036. Moreover, if we observe the projected distribution of female inhabitants by
age classes, there is a shift in the most expressive class, that is projected as being from 35 to 39 years
old in 2016 and, in 2036, is from 55 to 59 years old. Also, notice that this model projects an increase
in elderly people, and a decrease in younger people and in female births. This ageing is due to the
relatively low fertility rate, which translates into a smaller number of newborns relatively to the elderly.
Therefore, this effect is propagated along the years, leading to the ageing of female population.
Figure 6.5: Projected distribution of female inhabitants by age classes in Arroios using the model withoutmigrations in 2016, 2021, 2026, 2031 and 2036.
The model without migrations is clearly inadequate to a parish, since it lacks to explore migration
flows of its inhabitants, which, in a long term, results in large errors in projections.
6.2.2 Model with one migratory flow
Recall the model with one migratory flow detailed in 3.2, which considers an open system. The
values obtained for the Leslie matrix are the same as the ones presented in 6.2.1. The main difference
between this model and the one without migrations, is the added column vector multiplied by a factor,
representing the migration flow. This column vector is calculated based on 2001 and 2011 Census’ data,
and, therefore, in the following projections, it is assumed that its behaviour is constant along time. These
matrix’ values are given in the table below.
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flux0 flux1 flux2 flux3 flux4 flux5 flux6
Leslie Matrix I 3.63668 2.63716 0.0517544 0.143053 0.711082 0.878175 0.136603
flux7 flux8 flux9 flux10 flux11 flux12 flux13
Leslie Matrix I 0.055245 0.113591 0.0690748 0.00489464 -0.0414223 -0.084362 -0.118182
flux14 flux15 flux16 flux17 flux18 flux19 flux20
Leslie Matrix I -0.1638 -0.224038 -0.314274 -0.501457 -0.651249 -0.837236 -0.88391
Table 6.3: Proportion of the flux relatively to the estimated female residents in 2011 of the model withone migratory flow in Arroios obtained using 2001 and 2011 Census data.
If we plot 2011’s female population data and the projected female inhabitants using this model with
one migratory flow, we obtain an error of 6.14342 × 10−17, which doesn’t translate into changes in the
female’s number by age class, as seen in the graphic below.
Figure 6.6: Distribution of female inhabitants by age classes in Arroios using 2011’s Census data, andusing the model with one migratory flow.
Let us now analyse the obtained projections for 2016, 2021, 2026, 2031 and 2036.
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Figure 6.7: Projected total of female inhabitants in Arroios using the model with one migratory flow in2016, 2021, 2026, 2031 and 2036.
Figure 6.8: Projected distribution of female inhabitants by age classes in Arroios using the model withone migratory flow in 2016, 2021, 2026, 2031 and 2036.
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According to this model, the female population increased in 2006, reaching a total of 19784 women,
which was the maximum number of inhabitants predicted. Afterwards, in 2011 the female population suf-
fered a decrease of 12.61%, approximately, being followed by another increase, reaching 19062 women
in 2016. From this year, the number of female inhabitants is predicted to decrease until it reaches 12603
women, according to this model and maintaining the migratory flow, fertility and mortality rates of the
population.
In 2016, the projected population has 5 to 10 years-old as most expressive age class, while in 2021
the most expressive age class is projected as 40 to 44 years-old. Therefore, this model predicts the
ageing of the female population between these years, along with a decrease in the total number of
female inhabitants, contradicting the predicted increase tendency verified between 2011 and 2016. In
2026, the most predominant age class is expected to be from 45 to 49 years old, which corresponds to
the natural ageing of the most expressive age class in 2021. The same effect happens in 2031 and in
2036, where the most predominant age class is from 50 to 54 years-old and from 55 to 59 years old,
respectively. However, in 2036, the age class of 25 to 29 years old suffers a slight increase over 5 years,
almost reaching the same expressivity in female inhabitants as the most predominant age class.
The assumption of uniform distributed and steady immigration and emigration flows between 2001
and 2011, can be misleading in these projections, since a socioeconomic crisis occurred from 2010
implying more emigration compared to immigration. Therefore, it is being applied and amplified with
time with the usage of this model.
The model with one migratory flow can be improved to adjust more to the changes in immigration
and emigration flows verified with time.
6.2.3 Model with immigration and correction flows
The model detailed in 3.3 assesses a system open to migrations, considering it through two vectors:
proportion of the immigration vector, fluximm, and proportion of a correction vector that includes em-
igration and errors in estimation of immigration (it was based on Portugal’s data which can differ from
Arroios parish), fluxcor.
Again, the Leslie matrix has parameters presented in 6.2.1. The proportion of immigration and
correction flows were calculated based on 2001 and 2011 Census’ data, being presented in the following
tables. In these projections, it is assumed that their behaviour is constant and uniform with time. The
error obtained for this model, comparing with the observed female population in 2011, was 1.3486 ×
10−16.
Notice that the immigration over 85 years old was assumed to be 0, since the last age group consid-
ered in Portugal’s Census is 85 or more years old.
fluximm0 fluximm1 fluximm2 fluximm3 fluximm4 fluximm5 fluximm6
Leslie Matrix I 1.50613 1.16503 0.405383 1.05065 2.45095 1.41552 0.468952
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fluximm7 fluximm8 fluximm9 fluximm10 fluximm11 fluximm12 fluximm13
Leslie Matrix I 0.335111 0.214589 0.148746 0.108777 0.0991554 0.106146 0.0889245
fluximm14 fluximm15 fluximm16 fluximm17 fluximm18 fluximm19 fluximm20
Leslie Matrix I 0.0443614 0.0190279 0.00930098 0.00288786 0 0 0
Table 6.4: Proportion of the immigration flow relatively to the estimated female residents in 2011 of themodel with immigration and correction flows in Arroios obtained using 2001 and 2011 Census data.
fluxcor0 fluxcor1 fluxcor2 fluxcor3 fluxcor4 fluxcor5 fluxcor6
Leslie Matrix I 2.13055 1.47213 -0.353629 -0.907592 -1.73987 -0.537348 -0.332349
fluxcor7 fluxcor8 fluxcor9 fluxcor10 fluxcor11 fluxcor12 fluxcor13
Leslie Matrix I -0.279866 -0.100998 -0.079671 -0.103882 -0.140578 -0.190508 -0.207106
fluxcor14 fluxcor15 fluxcor16 fluxcor17 fluxcor18 fluxcor19 fluxcor20
Leslie Matrix I -0.208162 -0.243066 -0.323575 -0.504345 -0.651249 -0.837236 -0.88391
Table 6.5: Proportion of a correction flow that includes emigration and errors in estimation of immigrationrelatively to the estimated female residents in 2011 of the model with immigration and correction flowsin Arroios obtained using 2001 and 2011 Census data.
Notice that the total immigration of Arroios was obtained multiplying the proportion of Portugal’s
female external immigrants for each age class, by the number of female inhabitants in Arroios that didn’t
live in this parish five years ago. A more clear idea of how adjusted the estimation of the immigration of
Arroios between 2001 and 2011 can be given by the graphic below.
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Figure 6.9: Projected distribution of female inhabitants by age classes in Arroios using the model withone migratory flow in 2016, 2021, 2026, 2031 and 2036.
For example, the number of female babies entering Arroios surpasses the estimated value for the
observed age class’ immigration, since the correction factor is non-negative and even greater than the
estimated proportion for immigration. By the absolute minimum achieved between the age class of
20 to 24 years old in the proportion of the correction factor, we have two possible scenarios: whether
the population experienced strong immigration and emigration fluxes, prevailing a positive flux, or the
predicted immigration wasn’t as high as the observed one, possibly meaning that the immigration flows
in Arroios were less than the ones registered for Portugal. Notice that, after the age-class of 50 to 54
years-old, the emigration flow prevails over the immigration flow, since the correction entry is negative
and its absolute value is greater, which means that the older female classes are leaving Arroios.
This model reduces itself to the previous one (the correction flux is obtained by subtracting the esti-
mated immigration of Arroios to the estimated migratory balance) if the growth of the flows is maintained
constant with time, projecting the population the same way as pictured in Figures 6.6, 6.7 and 6.8.
However, contrarily to the model with one migratory flow in 3.2, we can project the population using
different coefficients α and β, since the immigration occurs at a different rate than the one observed for
the applied correction, and vice-versa.
We can fine-tune the projection of the female population by, instead of considering vectors of pro-
portions of immigration and correction flows, considering vectors of proportions of internal and external
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flows. This can be explained by the almost constant rate at which internal fluxes are detected and by
the fluctuations that the external flux is subject to, namely the socio-economic conditions of countries of
origin and destination.
6.2.4 Model with internal and external flows
Following the model with internal and external flows explained on 3.4, several projections were ob-
tained using a constant rate, an increase of 5% and a decrease of 5% in external rates, whilst the internal
rates were maintained constant with time.
The parameters of the Leslie matrix, closed for migrations, are the ones presented in 6.2.1. The
parameters for the internal and external flows are introduced below. The error obtained for this model
considering the observed female population in 2011 was 1.54418× 10−16.
fluxint0 fluxint1 fluxint2 fluxint3 fluxint4 fluxint5 fluxint6
Leslie Matrix I 3.51086 2.5375 0.0132182 0.0464248 0.493284 0.760772 0.0994145
fluxint7 fluxint8 fluxint9 fluxint10 fluxint11 fluxint12 fluxint13
Leslie Matrix I 0.0293956 0.101607 0.0628015 -0.0015631 -0.0494154 -0.0948082 -0.126845
fluxint14 fluxint15 fluxint16 fluxint17 fluxint18 fluxint19 fluxint20
Leslie Matrix I -0.168028 -0.225817 -0.315153 -0.501579 -0.65138 -0.837389 -0.884314
Table 6.6: Proportion of the internal flow relatively to the total female residents of the model with internaland external flows in Arroios obtained using 2001 and 2011 Census data.
fluxext0 fluxext1 fluxext2 fluxext3 fluxext4 fluxext5 fluxext6
Leslie Matrix I 0.125815 0.0996613 0.0385363 0.0966285 0.217798 0.117403 0.0371888
fluxext7 fluxext8 fluxext9 fluxext10 fluxext11 fluxext12 fluxext13
Leslie Matrix I 0.0258494 0.0119839 0.00627329 0.00645774 0.00799309 0.0104463 0.00866303
fluxext14 fluxext15 fluxext16 fluxext17 fluxext18 fluxext19 fluxext20
Leslie Matrix I 0.00422779 0.00177914 0.000878334 0.000122026 0.00013138 0.000153188 0.000403932
Table 6.7: Proportion of the external flow relatively to the total female residents of the model with internaland external flows in Arroios obtained using 2001 and 2011 Census data.
To achieve a better visualisation of the estimated internal and external flows verified between 2001
and 2011, the column vectors above were plotted as shown bellow.
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Figure 6.10: Estimated internal and external flows of female inhabitants by age classes in Arroios usingthe model with internal and external flows.
The movements between parishes to or from Arroios (internal flow) had more impact on the number
of female inhabitants than those coming or going to the outside of Portugal (external flow). This was
expected since it is easier to adapt to a new place of residence inside the same country (there are
almost no cultural clashes) than to a new country with a new cultural identity, for example.
The external flow has the age class between 20 and 24 years old as maximum, indicating that female
immigrants are more expressive in the total inhabitants in this age class. After the age class of 45 years-
old, the external flow remains positive, even though it doesn’t suffer significant changes in its value,
meaning that there were no significant immigration from or emigration to foreign countries. Curiously,
there is a small increase in female inhabitants coming from foreign countries in age classes between 55
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and 69 years old, which can be explained to the improved life and less expensive conditions Portugal
offers when compared with their country of origin.
The age class of 0 to 4 years old is more expressive in the total of female inhabitants in that age class
when we analyse the internal flow. Moreover, if we notice that the other local maximum occurred with 25
to 29 years old, we can assume that the mothers moved from other parishes in Portugal bringing small
children with them. From 55 years old, women started to move out of Arroios, probably to caregivers’
houses.
Notice that the internal flux suffers less fluctuations with time because the social and economic
conditions that affect the country are more or less similar for all parishes, and, usually, similar reasons
for leaving or arriving to a certain parish apply along time. However, this scenario of almost reaching
stability in flows does not occur in the case of external flows. Notice that there are a lot of social-
economic differences between countries with time, which makes the countries more or less attractive to
its possible inhabitants. Therefore, these flows are affected by crisis. For example, the social-economic
crisis felt in Portugal, that started in 2010, affected our results, since these flow calculations were based
in a period contemplating 2010 and 2011, where external immigration decreased and external emigration
increased.
Using a constant rate, the same projections of the model with one migratory flow, pictured in Figures
6.6, 6.7 and 6.8, are obtained.
If we project the female population with an increase of 5% and a decrease of 5% in the external flow,
maintaining the internal flow, the results do not differ much from the ones presented above, being the
reason why these are not displayed in this thesis.
6.2.5 Model with Leslie matrix that includes immigration and emigration I
If we apply the model with Leslie matrix that includes immigration and emigration I, detailed on 3.5,
we can obtain the Leslie matrix parameters below. To minimize the overall error between the calcu-
lated female population and the observed one in 2011, an iterative method was applied, and its output
corresponds to the parameters below.
F3 F4 F5 F6 F7 F8 F9
Leslie Matrix I 0.0648405 0.115457 0.177891 0.199013 0.0636024 0.0156161 0.000346013
Table 6.8: Fertility rates using the model with Leslie matrix that includes immigration and emigration IIand an iterative method.
P0 P1 P2 P3 P4 P5 P6
Leslie Matrix I 0.816699 1.18027 0.976328 1.75146 1.07149 1.05948 0.994028
P7 P8 P9 P10 P11 P12 P13
Leslie Matrix I 1.11693 0.952943 1.04718 0.907832 0.999886 0.869962 0.942388
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P14 P15 P16 P17 P18 P19
Leslie Matrix I 0.796705 0.808649 0.545391 0.506219 0.205634 0.24096
Table 6.9: Survival rates using the model with Leslie matrix that includes immigration and emigration IIand an iterative method.
The error for this Leslie matrix that includes migration, without performing any tuning, is 0.0101619,
and after using the iterative method is 0.00976734. Once more, notice that we are supposing that the
migrations are steady and uniform with time.
Figure 6.11: Distribution of female inhabitants by age classes in Arroios using 2011’s Census data, andusing the model with Leslie matrix that includes immigration and emigration I.
The predicted distribution obtained using the model in question is close between the age classes
of 40 to 84 years old and of more than 90 years old. The number of female inhabitants was more
undervalued between the age class of 10 to 14 years old, and more overvalued between 85 to 89 years
old. These shifts may occur because, when we integrate the internal and external flow vectors, referring
to 10 years, we must ”convert” them to characterise 5 years only, which is accomplished by simply
supposing that the female population follows an uniform distribution and, in that way, we only need to
divide the proportions by two.
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Next, we introduce the projections obtained for 2016, 2021, 2026, 2031 and 2036 using this Leslie
matrix.
Figure 6.12: Projected total of female inhabitants in Arroios using the model with Leslie matrix thatincludes immigration and emigration I in 2016, 2021, 2026, 2031 and 2036.
The figure above shows that this model with a Leslie matrix containing migrations predicts a signifi-
cant decrease in female population from 2001 to 2036, only having a slight increase between 2011 and
2016.
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Figure 6.13: Projected distribution of female inhabitants by age classes in Arroios using the model withLeslie matrix that includes immigration and emigration I in 2016, 2021, 2026, 2031 and 2036.
Moreover, if we observe the projected distribution of female inhabitants by age classes, there is a
shift in the most expressive class, that is projected as being from 35 to 39 years old in 2016 and, in 2036,
is from 55 to 59 years old. This model projects that the most predominant classes will tend to be those
between 20 to 70 years old, that is, there is no increase in the number of female children and elderly.
This might be due to the strong impact of immigration and emigration that occur in these age classes.
6.2.6 Model with Leslie matrix that includes immigration and emigration II
The model with Leslie matrix that includes immigration and emigration II, detailed on 3.6 is now
going to be applied in order to obtain projections for the female inhabitants. The coefficients of the
Leslie matrix presented below were obtained through the referred model and by applying an iterative
method that diminishes the overall error between the calculated female population and the observed
one in 2011.
F3 F4 F5 F6 F7 F8 F9
Leslie Matrix I 0.0608461 0.109119 0.168138 0.187409 0.0598817 0.0147132 0.000326433
Table 6.10: Fertility rates using the model with Leslie matrix that includes immigration and emigration IIand an iterative method.
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P0 P1 P2 P3 P4 P5 P6
Leslie Matrix I 0.868069 1.00968 1.03572 1.08096 1.36505 0.83779 1.07718
P7 P8 P9 P10 P11 P12 P13
Leslie Matrix I 1.03561 1.03418 0.969863 0.98868 0.924553 0.949489 0.870746
P14 P15 P16 P17 P18 P19
Leslie Matrix I 0.868008 0.751969 0.593521 0.4676 0.2264 0.225816
Table 6.11: Survival rates using the model with Leslie matrix that includes immigration and emigration IIand an iterative method.
The error for this Leslie matrix that includes migration, without performing any tuning, is 0.39237,
and after using the iterative method is 0.0526008. Once more, notice that we are supposing that the
migrations are steady and uniform with time.
Figure 6.14: Distribution of female inhabitants by age classes in Arroios using 2011’s Census data, andusing the model with Leslie matrix that includes immigration and emigration II.
The predicted distribution calculated using the model detailed above is very similar to the one ob-
served in 2011. Moreover, only the age classes of 0 to 4 years old, 10 to 14 years old and 15 to 19 years
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old, suffer some shift between those two distributions.
The projections obtained for 2016, 2021, 2026, 2031 and 2036 using this Leslie matrix are introduced
next.
Figure 6.15: Projected total of female inhabitants in Arroios using the model with Leslie matrix thatincludes immigration and emigration II in 2016, 2021, 2026, 2031 and 2036.
Contrarily to the models previously shown, the model with a Leslie matrix containing migrations II
projects the total number of female inhabitants with a slight decrease between 2001 and 2026, and,
from 2026 to 2036, a small increase is verified.
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Figure 6.16: Projected distribution of female inhabitants by age classes in Arroios using the model withLeslie matrix that includes immigration and emigration II in 2016, 2021, 2026, 2031 and 2036.
If we analyse the projected distribution of female inhabitants by age classes, we can observe that
the classes with more expression inside the female population are always between 20 to 69 years old,
corresponding to the age where ative inhabitants have. There is also a shift of the most expressive
classes between 2016 and 2036: in 2016, the most expressive class was 40 to 44 years old, while in
2036, it is expected to be 50 to 54 years old. This model projects a stable number of daughters and a
decrease on the number of elderly women. This model is very well adjusted to our case study, Arroios.
6.3 Partial Derivatives of Evolutionary Entropy
To achieve a better knowledge of the population and its models, some parameters of perturbation
to changes must be studied. In this section, we will explore, in the light of these parameters subject
to change, three Leslie matrices introduced before: without migrations 6.2.1, including immigration and
emigration I 6.2.5, and including immigration and emigration II 6.2.6.
6.3.1 Leslie matrix without migrations 6.2.1
The parameter r = log λ, the growth rate, satisfies a variational principle and, in this case, considering
a Leslie matrix without migrations,
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r ≈ log 0.734435 ≈ −0.308654. (6.9)
Therefore, we can reach to the conclusion that, if the population follows this model, it is decreasing
with time, with no sufficient renovation of the inhabitants to compensate the deaths.
The value of the reproductive potential is:
Φ = −0.497105. (6.10)
The evolutionary entropy has as value:
H = 0.188451. (6.11)
6.3.2 Leslie matrix that includes immigration and emigration I 6.2.5
The growth rate r = log λ of Leslie matrix that includes immigration and emigration I is:
r ≈ log 0.942799 ≈ −0.0589022. (6.12)
Even though the decline of the projected population is less abrupt than in the Leslie model without
migrations, there is still a decrease in inhabitants of this parish.
The value of the reproductive potential is:
Φ = −0.282154, (6.13)
which confirms the insufficient renovation of population, leading to a decrease in the total number of
inhabitants.
The evolutionary entropy has as value:
H = 0.223252. (6.14)
6.3.3 Leslie matrix that includes immigration and emigration II 6.2.6
The Leslie matrix that includes immigration and emigration II has as growth rate r = log λ:
r ≈ log 1.01356 ≈ 0.0134648. (6.15)
The value of the growth rate is positive, which means that the population is growing according to this
projection. However, it is not a significant increase in the number of inhabitants of this parish but rather a
stabilisation of it, even though it contradicts the population decline predicted in the previous Leslie matrix
models.
Nevertheless, the reproductive potential is negative:
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Φ = −0.210466. (6.16)
The evolutionary entropy has as value:
H = 0.22393. (6.17)
6.4 Population Sample from 2017 and the Application of Models
The relation of this thesis to a project called Conhecer Arroios, a partnership between Junta de
Freguesia de Arroios and Instituto Superior Tecnico de Lisboa, allowed us to obtain a population sample
of 2344 inhabitants. Notice that the total population in 2011 was 32262 inhabitants. The interviews
were made between 1st April 2017 and 13th May 2017. Therefore, these results will be compared to
the projections obtained for 2016. For the sake of the simplification, only female population will be
considered.
The sampling error, since we assume that the population is infinite, is given by:
E ≈ 1.29√2344
≈ 2.66%. (6.18)
The sampling method is further detailed on [5].
Comparing the projected distribution, for 2016, of the female population using models without migra-
tions and with one migratory flow and the observed female distribution, we reach the conclusion that
these are quite different.
The model without migrations predicts a population more mature and with less young people. On
the other hand, the model with one migratory flow overestimates the number of women between 10 to
14 years old, and underestimates the middle age class. However, neither of these two models correctly
project the most expressive age class, which was, according to the population sample retrieved in 2017,
from 40 to 44 years old.
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Figure 6.17: Projected distribution of female inhabitants using models without migration and with onemigratory flow in 2016, and distribution of female inhabitants of the population sample taken from 2017,by age classes in Arroios.
The projections for 2016 obtained using the models with Leslie matrix that includes immigration and
emigration I and II describe better the observed female distribution than using the two models previously
exploited.
The model with Leslie matrix that includes immigration and emigration I underestimates the number
of female inhabitants until the age class of 55 to 59 years old. Afterwards, the predicted number of
women is higher than the reality found in this sample. The most expressive age class in the female
sample was correctly predicted by the model with Leslie matrix that includes immigration and emigration
II. However, this model underestimates the youngest age classes and overestimates the eldest age
classes.
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Figure 6.18: Projected distribution of female inhabitants using models with Leslie matrix that includesimmigration and emigration I and II in 2016, and distribution of female inhabitants of the populationsample taken from 2017, by age classes in Arroios.
Since we assumed, in the construction of the models, that the migration flow was steady and it would
have the same behaviour of the one observed between 2001 and 2011, the comparison between our
projections and the real world results may lead us to the conclusion that more young couples, with and
without children, entered this parish than between the same period ten years before. Furthermore, the
emigration of people with more that 55 years old increased. Almost certainly, Arroios parish is even
more rejuvenated than our projections.
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Chapter 7
Conclusions
In this work, we explored several models and their characteristics, which were previously used by
Demetrius, Lewis, and Leslie, for example. The first two were already seen in literature: the basic model
without migratory flows, previously explored by Leslie; the model with one migratory flow, proposed by
Caswell. However, these do not include emigration, which is relevant to our real world application on the
Arroios parish. Therefore, some new models were explored: the model with immigration and correction
flows; the model with internal and external flows; the models with Leslie matrix that includes immigration
and emigration I and II. These last two do not use a conventional Leslie matrix such as the one originally
proposed by Leslie. Instead, those matrices integrate immigration and emigration flows, which is also a
new way, as far as we know, of modelling the population. This innovation allows us to check the impact
of migration in terms of partial derivatives of evolutionary entropy.
After, we derived and developed some measures to assess the changes to perturbations in micro-
scopic parameters that describe the system, such as evolutionary entropy, H, and generation time,
variational principles (growth rate, r, and reproductive potential, Φ), and robustness, R. Furthermore,
we developed the theoretical analysis of growth rate sensitivity. We also implemented some of these
measures in Mathematica, given the Leslie matrix as input. In the future, we would have wanted to
extend this implementation to other population matrices, such as Lefkovitch matrices.
Another new and original work, as far as we know, was the compression of the Leslie matrices, in
which we designed a method based on graphs that reduced the number of nodes by half with null error.
In a future work, we pretend to implement the proposed algorithm in languages such as R.
In this thesis we had the chance to apply and test all previously mentioned theoretical models and
measures in a real world context, namely with data retrieved concerning a parish in Lisbon called Arroios.
The fact that we applied our theoretical models to a real population generated very interesting remarks
such as the emigration of the elderly, due to the lack of nursing homes in this region, and high immigration
of children, due to the high number of young couples with children that were immigrants.
Our projections, in a general way, predict a decrease in the number of female inhabitants, which
is corroborated by measures such as the growth rate of the Leslie matrices. In fact, the Leslie matrix
without migrations has the smallest growth rate of the three studied, suggesting that the decrease in
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Arroios’ population is higher when only the inhabitants, without migrants, are considered. Furthermore,
this population without migrations has the lowest reproductive potential, which indicates a low capacity of
reproduction of the individuals and slow renovation of the intrinsic population with time. The evolutionary
entropy is also the lowest, suggesting that this population without migrations is more susceptible to
changes in microscopic parameters that describe the system, being more exposed to extinction. On
the other hand, the Leslie matrix that includes immigration and emigration II has the highest growth rate
and, even though it is close to zero, it is positive, leading to the conclusion that the population is slightly
growing but tending to stability. Another important measure of population growth is the reproductive
potential, which is the smallest but still negative, meaning that the renovation of individuals is not at a
rate that compensates the exits and deaths verified in this society with migrations. Also, the evolutionary
entropy value is the highest but yet corresponds to a population that is very vulnerable to perturbations.
Notice that the evolutionary entropy found for Arroios is low comparing to other endangered species. For
example, the Desert tortoise (Gopherus agassizii) has a higher evolutionary entropy value (H = 0.548)
[30].
Finally, we would like to apply our models and measures to other parishes and refine them with
more data concerning different years and, in that way, improve the robustness of our models and its
projections, and help the decision makers to know certain intrinsic aspects of a population.
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Bibliography
[1] A. Lotka and F. Sharpe. A Problem in Age-Distribution. Philosophical Magazine, 21(124), 1911.
[2] H. Bernardelli. Population Waves. Journal of the Burma Research Society, 31, 1941.
[3] E. Lewis. On the Generation and Growth of a Population. The Indian Journal of Statistics, 6:93–96,
1942.
[4] P. H. Leslie. On the Use of Matrices in Certain Population Mathematics. 1945.
[5] A. Ferreira, A. Martins, H. M. Oliveira, L. V. Melo, and R. Varela. Estudo da Dinamica Populacional
da Freguesia de Arroios. July 2017.
[6] G. Royle and E. W. Weisstein. Reducible Matrix.
[7] M. Boyle. Notes on the Perron-Frobenius Theory of Nonnegative Matrices.
[8] D. Serre. Matrices: Theory and Applications. Springer Science & Business Media, 2010.
[9] C. D. Meyer. Matrix Analysis and Applied Linear Algebra. SIAM, 2000.
[10] T. Edition, editor. Graph Theory. Graduate Texts in Mathematics. Springer, 2006.
[11] P. H. Leslie. Some Further Notes on the Use of Matrices in Population Mathematics. 1948.
[12] H. Caswell. Matrix Population Models: Construction, Analysis, and Interpretation. Sinauer Asso-
ciates, 2nd edition, 2001.
[13] L. P. Lefkovitch. The Study of Population Growth in Organisms Grouped by Stages. Biometrics, 21
(1):1–18, March 1965.
[14] J. F. Alves and H. M. Oliveira. Similarity of General Population Pseudo-Leslie Matrices. Applied
Mathematics & Information Sciences, September 2015.
[15] L. A. Demetrius and V. M. Gundlach. Directionality Theory and the Entropic Principle of Natural
Selection. 2014.
[16] M. N. Stephen J. Kirkland. Group Inverses of M-Matrices and Their Applications. Chapman and
Hall/CRC, 2012.
63
![Page 78: Structured Population Dynamics in a Real World Context · An algorithm that compresses Leslie matrices is further explained. Finally, this knowledge is applied in a real world context,](https://reader034.vdocuments.us/reader034/viewer/2022043019/5f3b1c520ebb2058554f39a8/html5/thumbnails/78.jpg)
[17] M. Williamson. Some Extensions of the Use of Matrices in Population Theory. Bulletin of Mathe-
matical Biophysics, (13-17), 21.
[18] M. Usher. A Matrix Approach of Renewable Resources, with Special Reference to Selection
Forests. Journal of Applied Ecology, 3:355–367, 1966.
[19] C. Bosch. Redwoods: a Population Model. Science, 172:345–349, 1971.
[20] O. Dieckmann and J. Metz. The Dynamics of Physiologically Structured Populations. Springer,
1986.
[21] H. Caswell. Matrix Models and Sensitivity Analysis of Populations Classified by Age and Stage: a
Vec-permutation Matrix Approach. 2011.
[22] A. Rogers. Multiregional Demography: Principles Methods and Extensions. John Wiley and Sons,
1995.
[23] H. Caswell and N. S. Gassen. The Sensitivity Analysis of Population Projections. Demographic
Research, 33(28):801–840, October 2015.
[24] H. Caswell. Matrix population models. Wiley StatsRef: Statistics Reference Online.
[25] W. Hamilton. The Moulding of Senescence by Natural Selection. Jornal of Theoretical Biology, 12
(1):12–45, September 1966.
[26] L. A. Demetrius. Growth Rate, Population Entropy, and Evolutionary Dynamics. Theoretical Popu-
lation Biology, 41(2):208–236, April 1992.
[27] L. A. Demetrius and S. Legendre. Evolutionary Entropy Predicts the Outcome of Selection: Com-
petition for Resources that Vary in Abundance and Diversity. Theoretical Population Biology, 83:
39–54, February 2013.
[28] L. A. Demetrius. Boltzmann, Darwin and Directionality Theory. Physical Reports, 530(1):1–85,
September 2013.
[29] H. Caswell. Matrix Population Models. Sinauer Associates, 2000.
[30] L. Demetrius and H. M. Oliveira. Studies on Evolutionary Entropy. Preprint, 2017.
[31] L. Demetrius, V. Gundlach, and M. Ziehe. Darwinian Fitness and the Intensity of Natural Selection:
Studies in Sensitivity Analysis. Journal of theoretical biology, 249(4):641–653, 2007.
[32] L. Arnold, L. Demetrius, and V. Gundlach. Evolutionary Formalism for Products of Positive Random
Matrices. The Annals of Applied Probability, 4(3):859–901, 1994.
[33] L. Demetrius, V. Gundlach, and G. Ochs. Complexity and Demographic Stability in Population
Models. Theoretical Population Biology, 65(3):211–653, 2004.
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Appendix A
Iterative Method
Function that receives the Leslie Matrix, the female population in 2001 and in 2011, the maximum
difference allowed between decrease the overall error, and the rational number to be added or subtracted
to each parameter until the model projects 2011 with a minimum error. Returns the matrix optimized
taking into account the error obtained using the data of 2011.
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iteradif = Function[{la, p2001, p2011, eps, step, indice}, Module[{dif, ma, pa,a, b, c, d, e, f, g, h, in, jn, l, m, n, o, p, q, r, s, t, u, el, ij, fi},
{a, b, c, d, e, f, g, h, in, jn, l, m, n, o, p, q, r, s, t, u} = PadRight[{}, 20];dif = Table[(MatrixPower[la, indice].p2001)[[i]] - p2011[[i]],
{i, 1, Dimensions[p2001][[1]]}];pa = {a, b, c, d, e, f, g, h, in, jn, l, m, n, o, p, q, r, s, t, u};ma =
Extract[Table[PadRight[PadLeft[{pa[[j]]}, j], Dimensions[p2001][[1]]], {i,1, Dimensions[p2001][[1]] - 1}, {j, 1, Dimensions[p2001][[1]] - 1}], {1}];
ma = Prepend[ma, PadRight[{}, Dimensions[p2001][[1]]]];ij = 2;While[ij ≤ Dimensions[p2001][[1]],If[Extract[dif[[ij]], {1}] != 0,el = Extract[dif[[ij]], {1}];While[Abs[el] > eps,If[Extract[dif[[ij]], {1}] > 0,
pa[[ij - 1]] = pa[[ij - 1]] + step;ma[[ij, ij - 1]] = pa[[ij - 1]];fi = la - ma;dif = Table[(MatrixPower[fi, indice].p2001)[[i]] - p2011[[i]],
{i, 1, Dimensions[p2001][[1]]}];el = Extract[dif[[ij]], {1}],
pa[[ij - 1]] = pa[[ij - 1]] - step;ma[[ij, ij - 1]] = pa[[ij - 1]];fi = la - ma;dif = Table[(MatrixPower[fi, indice].p2001)[[i]] - p2011[[i]],
{i, 1, Dimensions[p2001][[1]]}];el = dif[[ij]];
];];
];ij = ij + 1;
];Print[Extract[(Sum[Abs[((MatrixPower[fi, indice].p2001 - p2011) / p2011)[[i]]],
{i, 1, Dimensions[p2001][[1]]}]), {1}] / Dimensions[p2001][[1]]];Print[MatrixForm[fi]];fi
]
];
66