General Models:
Specific
Models: Examples in
text:
Directed Reciprocation
X
undertakes a significant cost to benefit Y, and Y in turn reciprocates that
benefit back to X. Operates
within or between species. (Trivers
1971) |
Partner
Fidelity Feedback: X and Y are associated
for an extended series of exchanges that last long enough that a feedback
operates: the effect of benefits transferred from X to Y returns and enhances
the fitness of X. Thus, by failing to cooperate, individual X ultimately
curtails its own fitness. (Bull and Rice 1991,
Nowak and May 1992, Frank 1994, Doebeli and Knowlton 1998, Simms and Taylor
2002 ) |
-Vertically
transmitted symbionts, optimal virulence evolution, ant-acacia mutualism. |
Partner
Choice: Either
individual X1 or X2 receives a benefit from Y,
depending on YÕs choice. Y chooses to interact with the X individual that
offers greater fitness returns. (Darwin 1859, Eshel
and Cavali-Sforza 1982, No‘ 1990, Bull and Rice 1991, Peck 1993, No‘ and
Hammerstein 1994, Batali and Kitcher 1995, Frank 1995, in press, Wilson and
Dugatkin 1997, West et al 2002b) |
-Squid-light organ
symbiosis, legume-rhizobium symbiosis, yucca-yucca moth symbiosis,
image-scoring in reef fish, allogrooming in impala. |
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Shared
Genes X
benefits X1 with which X shares alleles through descent from a
common ancestor. Operates
within species only. (Hamilton
1964a,b) |
Kin
Choice: X
recognizes
and directs benefits to more closely related X1 as opposed to more
distantly related X2 based on phenotype(s) of X1, X2. (Hamilton 1964a, Eshel
and Cavalli-Sforza 1982, Reeve 1989, Queller 2000) |
-Exclusion of
non-relatives in tunicates, GP9 locus in fire-ants, M-factors in beetles,
cell adhesion genes in social amoebae. |
Kin
Fidelity: X
directs benefits to X1 base upon X1Õs context dependent
spatial association with X. This proximity denotes shared genes with X. (Hamilton 1964a, Eshel
and Cavalli-Sforza 1982, Reeve 1989, West 2002a) |
-Parental
care in birds, coinfection in clonal microbes, aposemat-ism in familial
groups of aphids. |
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Byproducts X
benefits Y as a byproduct of an otherwise selfish act of X. Operates within or
between species. (West-Eberhard
1975, Brown 1983) |
One
Way: An act of X benefits Y as an automatic consequence
(byproduct) of XÕs self interested action. (West-Eberhard 1975,
Brown 1983, Connor 1995a) |
-Vultures and lions,
carrion feeders. |
Two
Way: Both
X and Y each benefit the other as automatic consequences (byproducts) of
their own selfish actions. Includes synergism: actions or coordinated
behavior that are automatically more fitness-enhancing when performed in
groups. (Hamilton 1971, Queller 1985, Connor
1995a) |
-Predator dilution in
bugs, selfish herds, Mullerian mimicry, unrelated helpers at the nest, ant
colonies founded my multiple queens. |
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Byproducts
Reciprocity: Y
evolves to enhance its benefit to X, which in turn increases the byproducts
it receives from X. The byproduct from X does not evolve, but the effect of Y
on X does.
(Connor
1986) |
-Honeyguide-man
mutualism. |