Objectives

By the end of this case study, you should be able to:

Explain what game theory is and how/why it is applied
to ecology.
Explain how novel behavioral strategies can invade a
population.
Explain evolutionarily stable strategy (ESS) and
frequency dependent selection.
Describe why natural selection acts on relative fitness
and how this fitness can depend on the frequency of
other "types” of individuals in the population.
Explain how coexistence of multiple
behavioural strategies in a population is
possible.
 me Theo
Game Theory is "a branch of
mathematics devoted to the study of
strategy in which players seek to
maximize their individual returns”.

’ w h o H=h Forms the basis of many economic theories,

war strategy, sociological theories (and
board games!)

John Maynard Smith emphasized the potential

for its use in behavioral ecology.

John Maynard 5mith

Game Theory
What behavioral strategy should individuals
adopt to maximize their individual fitness?
Does the “best” strategy depend on the
strategy of other individuals?

Should individuals behave aggressively and

fight for resources or behave non-
aggressively?
The i Ouesti n

If natural selection is “survival of the

I/ffesf,” then under what conditions
should individuals engage
aggressively in conflict for limited
resources? Can aggressive and non-
aggressive strategies coexist?
 Hawk vs. Dove Game
Two behavioral approaches
for
interactions over a contested resource
Hawks Doves
The les: Hawks
AGGRESSIVE
Always attack other individuals, taking the
resource if they win.
If a hawk encounters another hawk, it will win
half the time.
When it loses, it will suñer an injury cost.
The Rules: Doves
NON-AGGRESSIVE
When a dove encounters an opponent, it
may put on a aggressive display, but it does
NOT fight.
When a dove encounters a dove, it will win the
resource half the time.
When a dove encounters a hawk, the
hawk will always win the resource.
But since Doves don’t fight, they don’t incur
any injury costs.
Which behavioral strategy wins?

Hawks vs. Doves?

Is there an optimal behavioral strategy?
Will hawks take over?
Will doves be able to persist?
 Hawk Fitness

’! You are all HAWKS.

You will compete for the resource cards
which are surrogates for “reproductive
fitness”.
Play ROCK-PAPER-SCISSOR with your
neighbor. If a tie, repeat until someone
wins.
Winner gets 1 card from loser,
loser puts 1 card away
“permanently” (cost of fighting).
Repeat game 1 more time.
Typical Results

Figure I. Hawk Results

0

39
%

6 20M
Rese

Return cards so everyone has 4 resource cards again.

 Inerabil Invasion Hawks

Let’s see if a Dove (an individual with a random

mutation) can invade the population.

Mutant volunteer?
Vulnerability to Invasion - Hawks
Let’s see if a Dove (an individual with a
random mutation) can invade the
population.

Remember when a dove meets a hawk, the

dove always loses.

Watch what happens over 4 interactions with

our mutant Dove.
Reset

Return cards so everyone has 4 resource cards again.

Evolutionari Stable Strategy ESS)

ESS: “Behavioural strategy that is resistant

to invasion and most likely to be maintained
by natural selection”.

Is a Hawk an ESS? A Dove?

Dove Fitness
Now everyone are DOVES.
Play a ROCK-SCISSORS-PAPER game with
your neighbor.
Losers will give 1 card to the winners.
Doves do not suffer cost Scissors
so losers only lose that 1
card.
Repeat game 1 more time.
 Typical Results
2J I

Figure 2. Dove Results

0

1%

Z 279t›

3 4¥

Is this higher or lower than when all Hawks?

Typical Results
Figure 2. Dove Results
0

38°,

HIGHER! All hawks averaged 3 cards. Why?

Typical Results

Figure 2. Dove Results

No cost when losing as DOVE.

Reset

Return cards so everyone has 4 resource cards again.

 Inerabil Invasion Doves

Now let’s see if a HAWK can invade

a population of DOVES?

HAWK volunteer?
 Inerabil Invasion Doves

Now let’s see if a HAWK can invade

a population of DOVES?

What happens after 4 interactions?

Hawks vs. Doves: Evol
We are going to start with 10O o of the
population being Hawks and -90% Doves and
track what happens to this ratio over time.
Stand up and move around room, interacting with
as many others as you can.
Identify yourself as a HAWK or DOVE each time.
Remember:
Hawk-Hawk: Rock-Paper-Scissor & Winner gets 1 card
& loser pockets 1 card (cost)
Hawk-Dove: Hawk gets one card but Dove has no
cost. Dove-Dove: Rock-Paper-Scissor & Winner gets
1 card but loser has no cost.
Hawks vs. Doves
Hawks and Doves can reproduce their
own kind!

If you lose all your resource cards, you

are "dead” and must sit down.

If you get 8 resource cards, find someone

sitting down and convert them to whatever
you are (Hawk or Dove) by giving them 4
cards.
 Typical Results

• Hawks What was it Hawks

like early on? How 0.9 Doves

about later?
0.7

Dov
• Theory predicts for a ^.^
benefit to cost ratio of ‹
1:2, you would see
P op hon of
„ 50:50 proportion of
Hawks & Doves.
02

0.1

0 2 3 I
Time I ntervals
 1
Hawks
0.9
- W- Doves
0.8

0.7
Propor flon of Hawks and

0.6

05

0.0

0.3

0.2

01

0
0 1 2 3 4 5
Time Intervals
Frequency Dependent Selection

Frequency dependent selection occurs when the

fitness of an individual depends on the relative
frequency of other phenotypes in the population.
Positive frequency dependent selection — majority
advantage (majority phenotype wins).
Neqative frequency dependent selection —
minority advantage. Promotes genotypic and
phenotypic diversity. Some combination of
phenotype.
Frequencies may exist at which all phenotypes
have the same fitness.
Frequency Dependent Selecti n

When R = 50 and C = 100

When R = 50 and C = 75
SO 50
Doves Doves

20
10 10
o

-10 -10
•20 20
-30
-30
0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
Proportion Hawks Proporton Hawks

What are these graphs showing?

Left:Cost = 100 (higher)
Right: Cost =75 (lower)
Frequency Dependent Selecti n

When R = 50 and C = 100

When R = 50 and C = 75
50 SO

Doves Doves

30

20
10 10
o

-10 -10
•20 20
-30
-30
0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
Proportion Hawks Proporton Hawks

If relative cost hiqher, If relative cost is lower,

fewer Hawks. more Hawks.

If no cost?
Frequency Dependent Selecti n

When R = 50 and C = 100

When R = 50 and C = 75
50 SO

Doves Doves

30

20
10 10
o

-10 -10
•20 20
-30
-30
0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0
Proportion Hawks Proporton Hawks

If relative cost higher, If relative cost is lower,

fewer Hawks. more Hawks.

If no cost? All Hawks!

Frequency Dependent Selecti n

When R - 50 and C - 1N
When R - 60 and C - 76

F'roportbn Ha›d‹s
If Hawks start out lower than stable
equilibrium, relative fitness of Doves <
Hawks so Hawks increase in population
toward equilibrium.
Frequency Dependent Selecti n

When R = 50 and C = 100 When R = 50 and C = 75

Doves
Doves

0.5 80.9

If Hawks start out higher than stable

equilibrium, relative fitness of Doves > Hawks
so Hawks decrease in population toward
equilibrium.