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Topics in evolutionary dynamics
Lecture 1: Adaptation & the concept of fitness
François Massol
3rd summer school on Mathematical Biology
São Paulo, February 2014
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Lecture outline
1. Basics of evolution
2. Defining adaptation
3. The concept(s) of fitness
4. Fitness in different guises
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BASICS OF EVOLUTION
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Basics of evolution
What are the forces shaping the evolution of organisms?
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Basics of evolution
What are the forces shaping the evolution of organisms?
tim
e
population
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Basics of evolution
What are the forces shaping the evolution of organisms?
mutation
selection
migration
drift
tim
e
population
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Basics of evolution
What are the forces shaping the evolution of organisms?
mutation
selection
migration
drift tim
e
population
![Page 8: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/8.jpg)
Basics of evolution
What are the forces shaping the evolution of organisms?
mutation
selection
migration
drift tim
e
population
![Page 9: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/9.jpg)
Basics of evolution
What are the forces shaping the evolution of organisms?
mutation
selection
migration
drift tim
e
population
![Page 10: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/10.jpg)
Basics of evolution
What are the forces shaping the evolution of organisms?
mutation
selection
migration
drift tim
e
population
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Basics of evolution
What are the forces shaping the evolution of organisms?
mutation: new random variants
selection: heterogeneous degrees of success
migration: mixing with neighboring populations
drift: randomness takes its toll
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Basics of evolution
What are the forces shaping the evolution of organisms?
mutation: new random variants
selection: heterogeneous degrees of success
migration: mixing with neighboring populations
drift: randomness takes its toll
Evolution shapes the distribution of genotypes
Genotype = set of alleles carried at different loci
Genotype × Environment = Phenotype
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Basics of evolution
Population genetics formalism (p + q = 1)
Mutation
Selection
Migration
Drift
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Basics of evolution
Population genetics formalism (p + q = 1)
Mutation
Selection
Migration
Drift
1 1t A a t a A tp p q
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Basics of evolution
Population genetics formalism (p + q = 1)
Mutation
Selection
Migration
Drift
1 1t A a t a A tp p q
1 /t A tp w p w
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Basics of evolution
Population genetics formalism (p + q = 1)
Mutation
Selection
Migration
Drift
1 1t A a t a A tp p q
1 /t A tp w p w
1 1t t tp m p mP
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Basics of evolution
Population genetics formalism (p + q = 1)
Mutation
Selection
Migration
Drift
1 1t A a t a A tp p q
1 /t A tp w p w
1 1t t tp m p mP
1k N k
t t t
NkP p p q
kN
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Basics of evolution
Mutation / Selection / Migration / Drift
What else?
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Basics of evolution
Mutation / Selection / Migration / Drift
What else?
Homogamy vs. panmixia (mating system)
Dominance
Recombination
Epistasis (on selection)
Plasticity (environmental dependence)
Epigenetic effects
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Basics of evolution
Conditions for the evolution of traits
Differential success
Standing variance of trait values
Heritability of trait values
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Bergstrom & Dugatkin, “Evolution”
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DEFINING ADAPTATION
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Defining adaptation
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Defining adaptation
Evolution
Optimization
Natural selection
Adaptation
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Defining adaptation
Evolution
Optimization
Natural selection
Adaptation
process / history
outcome
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Defining adaptation
History-based definitions – Features built by natural selection for their current
role (Gould & Vrba)
– A derived character that evolved in response to a specific selective agent (Harvey & Pagel)
– Biological machinery or process shaped by natural selection to help solve one or more problems faced by the organism (Williams & Nesse)
– A is an adaptation for task T in population P iff A became prevalent in P because there was selection for A, where the selective advantage of A was due to the fact that A helped performed task T (Sober)
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Defining adaptation
History-based definitions – Features built by natural selection for their current
role (Gould & Vrba)
– A derived character that evolved in response to a specific selective agent (Harvey & Pagel)
– Biological machinery or process shaped by natural selection to help solve one or more problems faced by the organism (Williams & Nesse)
– A is an adaptation for task T in population P iff A became prevalent in P because there was selection for A, where the selective advantage of A was due to the fact that A helped performed task T (Sober)
Character arising through past natural selection
Problem: what character doesn’t?
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Defining adaptation
Nonhistorical definitions – A feature of the organism which interacts
operationally with some factor of its environment so that the individual survives and reproduces (Bock)
– Greater ecological-physiological efficiency than is achieved by other members of the populations (Mayr)
– An aspect of the developmental pattern which facilitates the survival and/or reproduction of its carrier in a certain succession of environments (Dobzhansky)
– A strategy that has the highest per capita growth given the conditions (Mitchell & Valone)
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Defining adaptation
Nonhistorical definitions – A feature of the organism which interacts
operationally with some factor of its environment so that the individual survives and reproduces (Bock)
– Greater ecological-physiological efficiency than is achieved by other members of the populations (Mayr)
– An aspect of the developmental pattern which facilitates the survival and/or reproduction of its carrier in a certain succession of environments (Dobzhansky)
– A strategy that has the highest per capita growth given the conditions (Mitchell & Valone)
Character conferring the best fitness among available ones
Minor problem: can this happen?
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Defining adaptation
A synthesis of definitions (Reeve & Sherman)
A phenotypic variant that results in the highest fitness among a specified set of variants in a given environment
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Defining adaptation
A synthesis of definitions (Reeve & Sherman)
A phenotypic variant that results in the highest fitness among a specified set of variants in a given environment
“optimization” in some sense
among available variants (contingent on history)
conditional on environmental state
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Defining adaptation
A synthesis of definitions (Reeve & Sherman)
A phenotypic variant that results in the highest fitness among a specified set of variants in a given environment
“optimization” in some sense
among available variants
conditional on environmental state
Pending question:
what is fitness?
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THE CONCEPT(S) OF FITNESS
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The concept(s) of fitness
The simplest: growth = fitness
( )dN
r N Ndt
Growth rate
1 ( )t t tN N N
Reproductive ratio
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The concept(s) of fitness
The simplest: growth = fitness
Problem: what if all genotypes grow or decline together?
distinguish absolute from relative growth rate
may depend on context / density / frequency
( )dN
r N Ndt
Growth rate
1 ( )t t tN N N
Reproductive ratio
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The concept(s) of fitness
The simple: selection = fitness
( )dN
r N Ndt
Relative growth rate
( )ii i
dfr N r f
dt
1 ( )t t tN N N
1
i t
it it
t
Nf f
N
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The concept(s) of fitness
What for?
time
frequency
1
3
2
4
1. invasibility
2. persistence
3. fixation
4. coexistence
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The concept(s) of fitness
What for?
time
frequency
1
3
2
4
1. invasibility
2. persistence
3. fixation
4. coexistence
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The concept(s) of fitness
What for?
time
frequency
1
3
2
4
1. invasibility
2. persistence
3. fixation
4. coexistence
Same problem: what happens with rare genotypes?
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The concept(s) of fitness
The simple: selection = fitness when rare
Invasion fitness
0
( )if
ir N r
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The concept(s) of fitness
The complex: unstable absence = fitness
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The concept(s) of fitness
The complex: unstable absence = fitness
0
( )if
ir N r
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The concept(s) of fitness
The complex: unstable absence = fitness
0
( )if
ir N r
( )ii i
dfr N r f
dt
![Page 44: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/44.jpg)
The concept(s) of fitness
The complex: unstable absence = fitness
0
( )if
ir N r
0
.
if
df
dtf
J
Jacobian matrix
/i j ijg f J i i ig r fr
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The concept(s) of fitness
The complex: unstable absence = fitness
More generally:
define invasion fitness as the leading eigenvalue of the Jacobian in the absence of the focal type
0
.
if
df
dtf
J
Jacobian matrix
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The concept(s) of fitness
Even more generally:
define invasion fitness as any criterion that qualifies whether the leading eigenvalue of the Jacobian in the absence of the focal type has negative real part
More than one fitness criterion might exist!
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FITNESS IN DIFFERENT GUISES
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Fitness in different guises
Population genetics fitness
With selection only
1
i t
it it
t
w Pp
wp
P
*
*
i
i
w P
w P
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Fitness in different guises
Population genetics fitness
Divergent selection in habitats x and y
Life cycle: Reproduction, regulation, total migration
Model = soft selection (Levene 1953)
iyixi x y
x y
w
w
wq
wq
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Fitness in different guises
Population genetics fitness
Divergent selection in habitats x and y
Life cycle: Reproduction, total migration, regulation
Model = hard selection (Dempster 1955; Ravigné et al. 2004)
x ix
y
iy
i
x x y
yq w q w
q w q w
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Fitness in different guises
Population genetics fitness
Divergent selection + environmental variability
Use matrices for reproduction, migration, ... events
…
Sp .ax . .m R E S D
Sp .ax . .m R E D S
Sp .max . E D S
Ravigné’s model
Levene’s model
Dempster’s model
Massol 2013
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Fitness in different guises
Stage-structured populations
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Fitness in different guises
Stage-structured populations
0 0 0 1 2 0 3
2 2
3
1 1 0
1 1
12 2
3 3
3
(1
(1 0 0
0 (1 0
0 0
)
)
)t t
s f
s
s f q s f s f s
q s q
q s q
q s s
s
N N
0 1 2 3
s0f0+ (1-q1)s1
q1s1 q2s2 q3s3
s∞ (1-q2)s2 (1-q3)s3
s0f3
s0f2 s0f1
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Fitness in different guises
Stage-structured populations
1t t N F G N
Fecundity + Survival/change in stage
0 0 0 1 2 0 3
2 2
3
1 1 0
1 1
12 2
3 3
3
(1
(1 0 0
0 (1 0
0 0
)
)
)t t
s f
s
s f q s f s f s
q s q
q s q
q s s
s
N N
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Fitness in different guises
Stage-structured populations
1t t N F G N
Fecundity + Survival/change in stage
0 1 2
0
3
0
0 0 0 0
0 0 0
0 0
0
0 0
Ts
ff f f
F e f
1 1
1 1
2
2
3
3 3
3
2
2
(1 0 0 0
(1 0 0
0
)
)
)(1 0
0 0
q
q s q
q s q
q
s
s
s
s
s
G 0 1,0,0,0e
0 0 1 2 3, , ,f f fs ff
![Page 56: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/56.jpg)
Fitness in different guises
Stage-structured populations
Formula for fitness (≠ maximal eigenvalue)
1t t N F G N
Fecundity + Survival/change in stage
1
0. .Tw f I G e 0 1,0,0,0e
0 0 1 2 3, , ,f f fs ff
0TF e f
![Page 57: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/57.jpg)
Fitness in different guises
Stage-structured populations
Formula for fitness (≠ maximal eigenvalue)
1t t N F G N
Fecundity + Survival/change in stage
Stage-wise fecundity
Time passed in each stage
1
0. .Tw f I G e 0 1,0,0,0e
0 0 1 2 3, , ,f f fs ff
0TF e f
![Page 58: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/58.jpg)
Fitness in different guises
Stage-structured populations
1t t N F G N
demonstration, french way
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Fitness in different guises
Stage-structured populations
1t t N F G N
1
( )
F G I I G I G F 1 I I
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Fitness in different guises
Stage-structured populations
1t t N F G N
1
Sp 1 Sp ( )
I G F 1 IF G
1
( )
F G I I G I G F 1 I I
![Page 61: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/61.jpg)
Fitness in different guises
Stage-structured populations
1t t N F G N
1
Sp 1 Sp ( )
I G F 1 IF G
1
( )
F G I I G I G F 1 I I
1( )max Sp
I G F 1 I
is a strictly decreasing function of λ
![Page 62: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/62.jpg)
Fitness in different guises
Stage-structured populations
1t t N F G N
1
Sp 1 Sp ( )
I G F 1 IF G
1
( )
F G I I G I G F 1 I I
1( )max Sp
I G F 1 I
is a strictly decreasing function of λ
1Sp max Sp 11
I G FF G
![Page 63: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/63.jpg)
Fitness in different guises
Stage-structured populations
1Sp max Sp 11
I G FF G
0TF e f
0 1,0,0,0e
0 0 1 2 3, , ,f f fs ff
![Page 64: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/64.jpg)
Fitness in different guises
Stage-structured populations
1Sp max Sp 11
I G FF G
0TF e f
0 1,0,0,0e
0 0 1 2 3, , ,f f fs ff
1 1
0Sp 0, T
I G F I G ef
Fitness! w
![Page 65: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/65.jpg)
Fitness in different guises
Stage-structured populations
A demographic re-interpretation
0TF e f
0 1,0,0,0e
0 0 1 2 3, , ,f f fs ff
1
0. .Tw f I G e
1
0 00
T T k
k
f I G e f G e
Offspring produced at time k
Lifetime offspring production
![Page 66: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/66.jpg)
Thank you for your attention!
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Further reading
Evolution, adaptation Grafen A. (1999). Formal Darwinism, the individual-as-maximizing-agent analogy
and bet-hedging. Proc R Soc Lond B Biol Sci, 266, 799-803.
Maynard Smith J. (1989). Evolutionary Genetics. Oxford University Press, Oxford.
Reeve H.K. & Sherman P.W. (1993). Adaptation and the goals of evolutionary research. Q Rev Biol, 1-32.
![Page 68: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/68.jpg)
Further reading
Fitness Caswell H. (2001). Matrix Population Models: Construction, Analysis, and
Interpretation, Second edition. 2nd edn. Sinauer Associates, Inc. Publishers, Sunderland, Massachussets.
Charlesworth B. (1994). Evolution in age-structured populations, Second edition. 2nd edn. Cambridge University Press, Cambridge.
Diekmann O., Heesterbeek J.A.P. & Metz J.A.J. (1990). On the definition and the computation of the basic reproduction ratio R0 in models for infectious diseases in heterogeneous populations. J Math Biol, 28, 365-382.
Roughgarden J. (1979). Theory of Population Genetics and Evolutionary Ecology: an Introduction. MacMillan publishing Co., Inc.
![Page 69: Topics in evolutionary dynamics - ICTP – SAIFR · Topics in evolutionary dynamics ... What are the forces shaping the evolution of organisms? ... Bergstrom & Dugatkin, “Evolution](https://reader033.vdocuments.us/reader033/viewer/2022051601/5ae2198b7f8b9a5b348bd4eb/html5/thumbnails/69.jpg)
Further reading
Selection & population genetics de Meeus T., Michalakis Y., Renaud F. & Olivieri I. (1993). Polymorphism in
heterogeneous environments, evolution of habitat selection and sympatric speciation: soft and hard selection models. Evol Ecol, 7, 175-198.
Débarre F. & Gandon S. (2011). Evolution in heterogeneous environments: between soft and hard selection. Am Nat, 177, E84-E97.
Ravigné V., Olivieri I. & Dieckmann U. (2004). Implications of habitat choice for protected polymorphisms. Evol Ecol Res, 6, 125-145.
Wallace B. (1975). Hard and soft selection revisited. Evolution, 29, 465-473.