1. Categories of Social Behavior
Actor
Cooperative behaviors can be favored when advantage to actor > receiver +
Selfish behaviors are always favorable
Selfish
Cooperative + -
Recipient
Spiteful
Altruism
Spiteful behaviors are theoretically possible if the harm to receiver >harm to actor
Altruistic behaviors are NEVER (by definition) favored through individual RS -

2. Evolution of Social Behavior
Altruism presents a paradox for natural selection:
If natural selection favors traits that increase individual fitness, how can we explain behaviors that cost an individual fitness while helping another?

3. Altruism Darwin hinted at an answer:
Selection could favor traits that result in a decrease of individual fitness if they INCREASE a RELATIVE’S fitness
r = coefficient of relatedness
r = the probability that two alleles in two individuals are identical by descent
By helping a relative--> more of “your” genes are passed to the next generation

4. Inclusive Fitness
An individual's fitness can be partitioned into direct and indirect components:
Inclusive fitness = Direct fitness + Indirect fitness
where
Direct W = personal reproductive success
Indirect W = RS of individuals that share alleles (weighted by the probability of sharing alleles, that is, relatedness)

5. Hamilton’s Rule Br> C
A gene for altruistic behavior would be favored by natural selection if:
Br> C
In other words, altruism may evolve when:
The cost to the actor (C) is low
The benefit to the recipient (B) is high
The action is between close relatives (high r)

6. Relatedness r relationship 1/2 parents-offspring; full-sibs
  1/4   grandparents-grandkids; half-sibs
  1/8   cousins (first cousins), uncle/aunt/niece/nephew
“I wouldn’t save a man from drowning but I would save two brothers or eight cousins,” - JBS Haldane

7. Haldane’s Rule predicts that Altruistic Behavior Should be Directed Toward Relatives
Belding’s Ground Squirrels (work by Paul Sherman)
Provide alarm calls to warn others of approaching predators.
Cost to actor -higher predation risk-->Benefit to receiver, reduced predation risk
males disperse from the natal territory, so that females in a colony tend to be closely related whereas males are unrelated to the females
Who screams?
Females more than expected
Males less than expected

8. When are alarm calls given?
Females are more likely to give alarm calls when close relatives are close by
Females are more likely to help close relatives chase away trespassers than nonrelatives
>

9. Kin Recognition
In order to direct altruistic behavior towards relatives, individuals must be able to recognize relatives

10. Cannibalistic Amphibians
David Pfennig polymorphic salamanders and tadpoles that occur in either omnivorous or carnivorous morphs
Cannibals actively avoid eating relatives (they suck them in, but spit them out). Experiments have shown that plugging the nares prevents recognition and cannibals will happily eat anyone.
Avoiding eating a relative improves one's inclusive fitness. Discrimination leads to > 2x siblings surviving at virtually 0 cost

11. Major histocompatibility Loci (MHC)
genes that code for membrane proteins that display antigens. It is thought that a greater diverstiy of MHC types allows more proteins to be recognized and therefore resistance to diseases is higher
both humans and mice avoid mating with individuals of similar MHC type. Mice can detect MHC similarity in the urine, while humans can (at least) detect it in sweat (The T-shirt box)
Female humans who are pregnant or taking oral contraceptives prefer males with similar MHC genotypes, Females not taking contraceptives prefer males with unrelated MHC genotypes - ie associate with relatives during child bearing and rearing, but not during mating
Recent evidence suggests perfume preferences are correlated with MHC genotypes - perfume might function to broadcast MHC type

12. Sooo MHC is a great example of how many processes work together:
sexual selection (MHC type affects mating success)
inbreeding (avoided to maintain high MHC diversity)
evolution of sex (maintains variance in MHC)
* coevolution with pathogens (avoiding specialization by one pathogen type)

13. Eusocilaity: “ultimate” example of altruism
True eusociality:
Overlapping generations
Cooperation among individuals in raising young
Specialized castes of individuals that are nonreproductive
Found in many insects (hymenoptera, termites, thrips), one group of mammals, and snapping shrimp
Reproductive female
Pheidole ants

14. Haplo-diploidy
Why would sterile castes give up all direct reproduction?
In Hymenoptera:
Males - 1n, develop from unfertilized eggs
Females - 2n develop from fertilized eggs 2n 1n
1/2
1/2 1
1/2 1 1n 2n 2n
females are more closely related to their sisters than their own offspring(3/4 vs 1/2, assuming the same dad)!
an allele spreads faster by helping mom reproduce than by reproducing itself!

15. But… • Some haplo-diploid species are not eusocial
• Some eusocial species are not haplo-diploid
Conclusion:
Haplo-diploidy may allow eusociality to evolve more easily, but it neither necessary nor sufficient for eusociality to evolve

16. Another case of Eusociality
Naked Mole Rats!
• Reproduction is by a single queen and 2-3 males
• Most matings are between parents & offspring or full-sibs--> r = 0.81
• Workers care for young, dig tunnels, defend colony
But workers would still be more related to their own offspring, so why don’t they reproduce?

17. Queens beat workers into submission
Naked Mole Rats
Queens beat workers into submission
• Queens shove non-relatives more than relatives
Work level
Before shove
After shove
All shoves
0.14
0.25
Tunnel shoves
0.34
0.58
Shoving by the queen increases effort by workers

18. Reciprocal Altruism
Can altruistic behavior to evolve even when directed to nonrelatives?
Yes!
Requires:
repeated interactions with other individuals
many opportunities (and an unpredictable number) to be altruistic
symmetrical costs and benefits among the interactants
When these conditions exist, what type of actions will natural selection favor?

19. Game Theory
Invented in 1940’s to analyze contrasting strategies in games (like poker, blackjack) --> later applied to economics, biology, etc.
Goal is to determine which strategy will give the largest average payoff over multiple repetitions

20. The Prisoner’s Dilemma
2 prisoners charged as accomplices are locked in separate cells
The punishment they receive depends on whether they cooperate with each other or defect and turn against each other
B’s action
Cooperate with A
Defect against A
A’s action
Cooperate with B
R (reward for cooperation)
S (sucker’s payoff)
Defects against B
T (temptation to cheat)
P (punishment for mutual defection)

21. The Payoff B’s action Cooperate with A Defect against A A’s action
Cooperate with B
R--> A gets 3
B gets 3
S--> A gets 0
B gets 5
Defects against B
T --> A gets 5
B gets 0
P--> A gets 1
B gets 1
For individual A: T>R>P>S and R>(S+T)/2
For one play, the highest payoff for A would be T
But, if they play again, what’s the probability B will play the sucker again?

22. Over the long term, what’s the best strategy?
Can be shown through economic analysis (game theory)
“tit-for-tat” = an individual starts by cooperating and then simply does whatever the opponent did in the previous round
This is an ESS (evolutionary stable strategy) --> cannot be invaded by mutant strategy
Another? “pavlov” = win-stay, lose-switch
Play on-line for yourself!

23. Vampire Bats
Vampires - forage at night for blood meals on large mammals
33% of young bats and 7% of adults fail to feed on any given night - 3 consecutive bloodless nights and a bat dies
vampires roost in small groups, and membership in groups changes, some members are regular associates and others aren’t
vampires will share bloodmeals each other, preferentially to related bats, but also to those with whom they have some experience

24. Bloodmeal Sharing in Vampire Bats
Bats preferentially share with nonrelatives that they are frequent roostmates with
Bats preferentially share with relatives, especially those related by more than 1/4