You are on page 1of 49

Cooperation and mutualism

Termite with digestive protozoan symbionts


ca 100 million years old
Cooperation
A behaviour which provides a benefit for another individual (recipient), and
which has been selected for because of its beneficial effects on the other
recipient
West et al. 2006. J Evol Biol 20:415-432

Hamiltons categories of social interaction


Effect on Recipients LIFETIME fitness

+ -

Effect on + Mutualism Selfishness


Actors
LIFETIME
fitness - Altruism Spite
Cooperation
A behaviour which provides a benefit for another individual (recipient), and
which has been selected for because of its beneficial effects on the other
recipient
West et al. 2006. J Evol Biol 20:415-432

Not include behaviour which has benefit


to others as a byproduct

e.g. Zebra fleeing lion alerts others to


danger, but this not cooperation

e.g. Elephant dung provides food for dung beetle larvae


but this not cooperation
Cooperation among relatives
Cooperation among non-relatives

A. Intraspecific cooperation among non-relatives


Unrelated helpers in cooperative breeders
Food sharing
Alarm calling
Coalitions
Allogrooming

B. Interspecific mutualisms
Cleaning relationships
Protection-provisioning
Insect-plant mutualisms
A. Intraspecific cooperation among non-relatives

Unrelated helpers in cooperative breeders

Pied kingfisher
When food scarce, groups accept secondary helper
(unrelated) (Reyer 1984; see DKW)
Secondary helper often replace primary breeder in
following season

Paper wasps, Polistes dominulus


20-30% of helpers are non-relatives
May inherit nest later
A. Intraspecific cooperation among non-relatives

Food sharing

Vampire bats: regurgitate blood meals


Starve in 60 hours without blood

Chimpanzees: share food with non-relatives


(especially meat)
A. Intraspecific cooperation among non-relatives

Alarm calling

Many species of birds give alarm call which


benefits other individuals

e.g. White browed scrub wren


Leavesley & Magrath 2005. Anim Behav
70:365-373
A. Intraspecific cooperation among non-relatives

Coalition-formation and allogrooming

Olive baboons (Packer 1977; see KDW)


Males enlist the support of other males to gain
access to oestrus females
Males tended to request aid from an idndividual
who in turn requested aid from him.

Vervet monkeys (Cheney & Seyfarth 1984)


Grooming relationships among non-relatives
Individuals respond to playbacks of another if
recently groomed by them
B. Interspecific mutualisms

Cleaning relationships

e.g. cleaner fish client


Cleaner remove parasites
Also cheat by feeding on mucus, scales

e.g. Banded mongoose - warthog


B. Interspecific mutualisms

Protection-provisioning
e.g. Ant-lycaenid butterfly larvae
Larvae of butterflies of family Lycaenidae
secrete substances which attract or reward ants
Ants protect larvae from parasitic wasps and fly
predators

6000 species Lycaenid butterfly, 75% associated with


ants
Wide range of mutual dependency: obligate to
facultative to parasitic
B. Interspecific mutualisms

Mutual provisioning
e.g. Honeyguide-human
Honeyguides guide humans to bee nests
- Humans destroy the nest for the honey, bird benefits
from leftovers (eats eggs, larvae and beeswax)
Boran tribe (N. Kenya):

Ed Yong Not exactly rocket science Blog


B. Interspecific mutualisms

Insect-plant mutualisms
e.g. Ant-acacia
Acacia produce ant houses (domatia)
Ants protect against browsers

e.g. Fig wasps fig


Fig provide nest site, fig wasp pollinate fig

e.g. Flowers and pollinators


Flowers provide nectar, insects pollinate flowers
How does cooperation evolve?

1. Kin selection Helping relatives


2. Mutualism Win-win
3. Enforcement Cooperate or else
4. Reciprocity You scratch my back
2. Mutualism

Cooperation most easy to explain if fitness effects are (+/+)

e.g. Mongoose - warthog

Both benefit immediately from the mutualism


Mutualism

Cooperation most easy to explain if fitness effects are (+/+)


e.g. cooperative hunting in lions (also African wild dogs)

Single lioness not very effective at hunting


Prey capture maximized when mutliple females hunt together
Benefits of increased success rate (and larger prey) outweigh costs of sharing
meat (Caraco & Wolf 1975; Stander 1992 Behav Ecol Sociobiol 29:445-454)
Mutualism

Cooperation most easy to explain if fitness effects are (+/+)

Sometimes actions are self-interested, but benefits are deferred

Olive baboon Pied kingfisher Long-tailed manakin


Mutualism: deferred benefits
Group augmentation (Kokko et al 2001; PRSL):
Proposed to explain unrelated helpers in many
cooperative breeders

Helpers help so that inherit a large productive


group in future
Meerkats

Nest inheritance explains the presence of


unrelated foundresses and why they help
(Leadbeater et al. 2011 Science 333:874-876)

Polistes dominulus
3. Enforcement / manipulation
Cooperation may be due to deception or coercion

e.g. Cuckoo chick

e.g. Cooperative cichlids: pay to stay

e.g. Fish size hierarchies: threat of eviction

Buston 2003. Nature 424:145-146


4. Reciprocity
Cooperation can pay if the recipient pays back later
i.e. the deferred benefits are due to an altruistic response later, by the
recipient
Reciprocal altruism (Trivers 1972)

Actor pays small cost


Recipient gets large benefit

Later

Recipient repays actor

Both benefit
4. Reciprocity
Cooperation can pay if the recipient pays back later
i.e. the deferred benefits are due to an altruistic response later, by the
recipient
Reciprocal altruism (Trivers 1972)

Problem is that there is


temptation to cheat

Individuals who take advantage


but dont give back their fair share
Prisoners dilemma
Captures the problem in most basic form
Gain by cooperating, gain most by cheating a cooperator

Player 2

Cooperate Defect

3 5
Cooperate
3 0
Player 1
0 1
Defect
5 1
Prisoners dilemma
Captures the problem in most basic form
Gain by cooperating, gain most by cheating a cooperator

Player 2

Cooperate Defect

3 5
Cooperate
3 0
Player 1
0 1
Defect
5 1
Prisoners dilemma
Captures the problem in most basic form
Gain by cooperating, gain most by cheating a cooperator

Player 2

Cooperate Defect

3 5
Cooperate
3 0
Player 1
0 1
Defect
5 1
Cooperation in the Prisoners dilemma

Two main factors promote cooperation


1. Repeated interactions:
- increases cost of defection (lose out on long-term success)

2. Conditional behaviour
- cooperation is conditional upon the other partners behaviour

Robert Axelrod: Held a computer tournament (1979) where contestants entered a


strategy, e.g. Cooperate for X rounds, then defect once

Winner =Tit for Tat


Tit for Tat

Cooperate on first round, then do whatever other player did on last round.

Tit for Tat C C C C D D C C


Opponent C C C D D C C C

Tit for tat successful because


1. Nice
2. Provocable
3. Forgiving
4. Clear and simple
Tit for tat in humans

First world war (1914-18)


Live and let live system

I was having tea with A Company when we heard a lot of shouting and went out to
investigate. We found our men and the Germans standing on their respective parapets.
Suddenly a salvo arrived but did no damage. Naturally both sides got down and our men
started swearing at the Germans, when all at once a brave German got on to his parapet
and shouted out "We are very sorry about that; we hope no one was hurt. It is not our
fault, it is that damned Prussian artillery." (Rutter 1934, p. 29)

See Axelrod, R. (1984). The evolution of cooperation. Penguin Science Press


Do animals play tit for tat?
Repeated interactions do promote cooperation
e.g. Cleaner fish only occur on coral reefs, not in open seas
- Cleaners more cooperative with more faithful clients

But do animals employ tit for tat strategy?

Milinksi (1987) tested using sticklebacks in a tank


- Sticklebacks inspect predators: gain information on exact position,
motivation, identity of predator
- Often inspect in pairs
cm = Cooperative mirror: When move forward, other player move forward
dm = Defecting mirror: When move forward, other player appear to move away

Results
Stickleback move closer to predator when with a cooperative player
Milinksi 1987. Nature 325:433-435
Convincing?
Stephens et al. 1997 repeated the experiment but with the proper control:
an opaque partition so predator was no longer visible (Animal Behaviour
53:113-133)

Result: The ability to see the predator had no effect on the median
proximity to the predator.

Behaviour that was supposedly tit for tat arise because of two effects:
sticklebacks like to approach predators AND other sticklebacks
In general, little evidence of tit for tat in animals
Most examples of cooperation between non-relatives involve an immediate
benefit or manipulation (Clutton-Brock 2009 Nature 462: 51-57)

Recent syntheses (Sachs et al 2004; Foster & Wenseleers 2006) focus on


three processes that can enhance the benefits of cooperation

1. Partner fidelity (i.e. high chance of repeated interaction)


2. Partner feedback (investment in cooperation helps other partner to invest
in cooperation)
3. Partner choice and sanctions (leave non-cooperative individuals)

Sachs et al 2004 Q. Rev Biol. 79:135-160


Foster & Wenseleers 2006 J Evol Biol 19:1283-1293
Human cooperation

Human cooperation unusual in a number of respects:


Cooperate in large groups
Cooperate with non-relatives
Cooperate in one-shot scenarios (e.g. restaurant tipping)
Possess inherent sense of fairness

Henrich et al 2001: Ultimatum game in 15 small scale societies


Behavioral and Brain Sciences 28:795-855

Levels of cooperation and generosity correlated with dependency on


cooperation and trade
A cooperative brain?
Rule: A card with a vowel on the front always has an even number
on the back

A H 4 7

Q: Which card or cards should you definitely turn over to see if


the rule has been broken?
Results

?
A cooperative brain?
Rule: anyone drinking beer must be at least 21 years of age

Drinking Drinking
25 16
Beer Coke years years

Card have drink on one side, age of drinker on the other


Q: which card or cards do you need to turn over to check
whether anyone in the bar is breaking the law?
The Wason Test

The two tests are logically identical A H 4 7


[A,H,4,7] and [Beer, Coke, 25, 16]

<25% get it right on the first version 25 16


Beer Coke
years years
>75% get it right on the second version

Cosmides & Tooby (1992) argue that we have an inbuilt cheat detection
module of the brain (The Adapted Mind, OUP)

Helps us to detect free riders quickly


Self deception
Cheaters can prosper if they can avoid detection, but our brains are wired up to
spot them (Wason test)

But deception possible if deceive yourself!

Self-deception: Hiding the truth from yourself to hide it more deeply from
others. (Trivers 1991)

Trivers RL 1991. Ann New York Acad Sci 907:114-137


Trivers RL 2011. Deceit and Self-Deception. Allen Lane
Adams et al. 1996 J. Abn. Psych. 105:440
Test subjects given a questionnaire, divided into homophobic and non-
homophobic groups

The shown sexual videos: heterosexual, lesbian, homosexual sex


Plethysmograph around the base of the penis measured changes in circumference
Heterosexual video

Adams et al. 1996 J. Abn. Psych. 105:440

Results: Lesbian video

1. Only homophobic men aroused by


homosexual video
2. All men give accurate estimates of
tumescence with one exception:

Homophobic men deny their response to the


male homosexual video Homosexual video

homophobic

non-homophobic
Emotions
Frank (1987) Passions within Reason: the strategic role of the emotions
Emotions (shame, guilt, love) are an adaptation: Serve a dual purpose
1. Reward / Punish cooperative or non-cooperative behaviour (satisfaction,
outrage, guilt, shame)
2. Act as honest signals of cooperative nature
Hard to fake e.g. Blushing, body language when lying

Lack of emotion is a detectable indicator of non-cooperative


nature

Cleckley 1941 The Mask of Sanity


Psychopath is a perfect mimic of normal functioning
person

Ronson J (2012) The Psychopath Test


Love
Humans behave irrationally in many ways
Frank (1987) argues that irrational emotions may be adaptive, help to solve
commitment problems
Love
Humans behave irrationally in many ways
Frank (1987) argues that irrational emotions may be adaptive, help to solve
commitment problems
An example: people fall in love
Searching for a partner is costly, so it is rational to settle on a partner before
having examined all potential candidates
But after a partner is chosen, circumstances may change (e.g. a more
attractive partner may come along)
Love allows humans to commit to long term cooperation:
Promotes fidelity, teamwork

Love is not love


Which alters when it alteration finds
Or bends with the remover to remove;
O, no! it is an ever-fixed mark,
That looks on tempests and is never
shaken William Shakespeare, Sonnet 116
Empathy and Fairness

Franz de Waal: Chimps, other primates have a sense of


fairness

E.g. Capuchin experiment (Brosnan & de Waal 2003 Nature)


Brosnan & de Waal 2003.
Nature 425, 297
Empathy
Empathy defined as The ability to understand and share the
feelings of others

Why did empathy evolve?

Human evolution is characterised by a series of expansions of


sphere of empathy:
Family clan tribe nation- .??
We are the Borg. Your biological and
technological distinctiveness will be
added to our own. Resistance is futile.
Key points

Cooperation among non-relatives is common in nature

Explanations based on mutualism, manipulation, reciprocity

Mutualism and manipulation common in animals

Reciprocity is fundamental to human cooperation

Human brains possess mechanisms to enforce honesty and detect


cheats

Human cooperation (and possibly some animals) characterised by


empathy and sense of fairness