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

Hamilton’s categories of social interaction
Effect on Recipient’s LIFETIME fitness

+ -

Effect on + Mutualism Selfishness
Actor’s
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 .

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 . Intraspecific cooperation among non-relatives •  Unrelated helpers in cooperative breeders Pied kingfisher When food scarce. A. groups accept secondary helper (unrelated) (Reyer 1984.

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. Anim Behav 70:365-373 . White browed scrub wren Leavesley & Magrath 2005.g. Intraspecific cooperation among non-relatives •  Alarm calling Many species of birds give alarm call which benefits other individuals e.

” Vervet monkeys (Cheney & Seyfarth 1984) Grooming relationships among non-relatives Individuals respond to playbacks of another if recently groomed by them . 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. Intraspecific cooperation among non-relatives •  Coalition-formation and allogrooming Olive baboons (Packer 1977. A.

g. scales e. cleaner fish – client Cleaner remove parasites Also cheat by feeding on mucus. Banded mongoose . Interspecific mutualisms •  Cleaning relationships e.warthog .g. B.

Interspecific mutualisms •  Protection-provisioning e. B. 75% associated with ants Wide range of mutual dependency: obligate to facultative to parasitic .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.

Humans destroy the nest for the honey. Honeyguide-human Honeyguides guide humans to bee nests . larvae and beeswax) Boran tribe (N.g. bird benefits from leftovers (eats eggs. Kenya): Ed Yong ‘Not exactly rocket science’ Blog . B. Interspecific mutualisms •  Mutual provisioning e.

Flowers and pollinators Flowers provide nectar. B.g.g. Fig wasps – fig Fig provide nest site. fig wasp pollinate fig e. Ant-acacia Acacia produce ‘ant houses’ (domatia) Ants protect against browsers e. insects pollinate flowers .g. Interspecific mutualisms •  Insect-plant mutualisms e.

Reciprocity ‘You scratch my back…’ .  Mutualism ‘Win-win’ 3.  Enforcement ‘Cooperate or else’ 4. Kin selection ‘Helping relatives’ 2. How does cooperation evolve? 1.

2.warthog Both benefit immediately from the mutualism . Mutualism •  Cooperation most easy to explain if fitness effects are (+/+) e. Mongoose .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 (+/+) e.g.

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. 2011 Science 333:874-876) Polistes dominulus . 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.

Nature 424:145-146 . Cuckoo chick e. Cooperative cichlids: ‘pay to stay’ e. 3.g.g.g. Fish size hierarchies: threat of eviction Buston 2003. Enforcement / manipulation •  Cooperation may be due to deception or coercion e.

4. the deferred benefits are due to an altruistic response later.e. Reciprocity Cooperation can pay if the recipient pays back later i. by the recipient Reciprocal altruism (Trivers 1972) Actor pays small cost Recipient gets large benefit Later… Recipient repays actor …Both benefit .

4. 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 don’t give back their fair share .e. Reciprocity Cooperation can pay if the recipient pays back later i.

Prisoner’s 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 .

Prisoner’s 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 .

Prisoner’s 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 .

e. Cooperation in the Prisoner’s dilemma Two main factors promote cooperation 1.cooperation is conditional upon the other partner’s behaviour Robert Axelrod: Held a computer tournament (1979) where contestants entered a strategy. then defect once’ Winner =Tit for Tat .g.  Repeated interactions: .  Conditional behaviour . ‘Cooperate for X rounds.increases cost of defection (lose out on long-term success) 2.

‘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. Tit for Tat Cooperate on first round. then do whatever other player did on last round.  Provocable 3.  Clear and simple .  Forgiving 4.  Nice 2.

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. (1984). It is not our fault. Penguin Science Press . We found our men and the Germans standing on their respective parapets. R. Naturally both sides got down and our men started swearing at the Germans. 29) See Axelrod. The evolution of cooperation. we hope no one was hurt." (Rutter 1934. p. Suddenly a salvo arrived but did no damage. it is that damned Prussian artillery. when all at once a brave German got on to his parapet and shouted out "We are very sorry about that.

Often inspect in pairs .Cleaners more cooperative with more faithful clients But do animals employ tit for tat strategy? Milinksi (1987) tested using sticklebacks in a tank . Cleaner fish only occur on coral reefs.Sticklebacks ‘inspect’ predators: gain information on exact position. not in open seas . motivation. Do animals play tit for tat? •  Repeated interactions do promote cooperation e.g. identity of predator .

‘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 .cm = Cooperative mirror: When move forward.

Nature 325:433-435 .Milinksi 1987.

’ Behaviour that was supposedly tit for tat arise because of two effects: sticklebacks like to approach predators AND other sticklebacks . 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.

  Partner choice and sanctions (leave non-cooperative individuals) Sachs et al 2004 Q. Foster & Wenseleers 2006) focus on three processes that can enhance the benefits of cooperation 1. 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.  Partner fidelity (i.e. high chance of repeated interaction) 2. In general.  Partner feedback (investment in cooperation helps other partner to invest in cooperation) 3. 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 ? .

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? . 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.

7] and [Beer. Coke. The Wason Test •  The two tests are logically identical A H 4 7 [A.H.4. 25. OUP) •  Helps us to detect free riders quickly . 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.

Deceit and Self-Deception.“ (Trivers 1991) Trivers RL 1991. 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. Ann New York Acad Sci 907:114-137 Trivers RL 2011. Allen Lane . Self deception Cheaters can prosper if they can avoid detection.

divided into homophobic and non- homophobic groups •  The shown sexual videos: heterosexual. homosexual sex •  Plethysmograph around the base of the penis measured changes in circumference . Psych. lesbian. 105:440 Test subjects given a questionnaire. Abn.Adams et al. 1996 J.

Psych. Heterosexual video Adams et al. 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 . Abn. Only homophobic men aroused by homosexual video 2. 1996 J. 105:440 Results: Lesbian video 1.

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

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. 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. Love Humans behave irrationally in many ways Frank (1987) argues that irrational emotions may be adaptive. That looks on tempests and is never shaken William Shakespeare. so it is rational to settle on a partner before having examined all potential candidates But after a partner is chosen.g. O. no! it is an ever-fixed mark. circumstances may change (e. Sonnet 116 .

other primates have a ‘sense of fairness’ •  E.g. Empathy and Fairness •  Franz de Waal: Chimps. Capuchin experiment (Brosnan & de Waal 2003 Nature) .

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Nature 425.Brosnan & de Waal 2003. 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. Resistance is futile.” . Your biological and technological distinctiveness will be added to our own.

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 .