Frequency-dependent selection

Frequency-dependent selection is the term given to an evolutionary process by which the fitness of a phenotype depends on its frequency relative to other phenotypes in a given population.

• In positive frequency-dependent selection, the fitness of a phenotype increases as it becomes more common.
• In negative frequency-dependent selection, the fitness of a phenotype decreases as it becomes more common. This is an example of balancing selection.

Frequency-dependent selection is usually the result of interactions between species (predation, parasitism, or competition), or between genotypes within species (usually competitive or symbiotic), and has been especially frequently discussed with relation to anti-predator adaptations. Frequency-dependent selection can lead to polymorphic equilibria, which result from interactions among genotypes within species, in the same way that multi-species equilibria require interactions between species in competition (e.g. where α_ij parameters in Lotka-Volterra competition equations are non-zero).

Negative frequency-dependent selection

Anvil stone, where a thrush has broken open the shells of polymorphic grove snails, may be a sign of frequency-dependent selection.

The first explicit statement of frequency-dependent selection appears to have been by Edward Bagnall Poulton in 1884, on the way that predators could maintain color polymorphisms in their prey.

Perhaps the best known early modern statement of the principle is Bryan Clarke's 1962 paper on apostatic selection (a synonym of negative frequency-dependent selection). Clarke discussed predator attacks on polymorphic British snails, citing Luuk Tinbergen's classic work on searching images as support that predators such as birds tended to specialize on common forms of palatable species. Clarke later argued that frequency-dependent balancing selection could explain molecular polymorphisms (often in the absence of heterosis) in opposition to the neutral theory of molecular evolution.

Another example is plant self-incompatibility alleles. When two plants share the same incompatibility allele, they are unable to mate. Thus, a plant with a new (and therefore, rare) allele has more success at mating, and its allele spreads quickly through the population .

In human pathogens, such as the flu virus, once a particular strain has become common, most individuals have developed an immune response to that strain. But a rare, novel strain of the flu virus is able to spread quickly to almost any individual, causing continual evolution of viral strains.

The major histocompatibility complex (MHC) is involved in the recognition of foreign antigens and cells. Frequency-dependent selection may explain the high degree of polymorphism in the MHC.

In behavioral ecology, negative frequency-dependent selection often maintains multiple behavioral strategies within a species. A classic example is the Hawk-Dove model of interactions among individuals in a population. In a population with two traits A and B, being one form is better when most members are the other form. As another example, male common side-blotched lizards have three morphs, which either defend large territories and maintain large harems of females, defend smaller territories and keep one female, or mimic females in order to sneak matings from the other two morphs. These three morphs participate in a rock paper scissors sort of interaction such that no one morph completely outcompetes the other two. Another example occurs in the scaly-breasted munia, where certain individuals become scroungers and others become producers.

Positive frequency-dependent selection

Venomous coral snake's warning coloration can benefit harmless mimics, depending on their relative frequency.

Where negative frequency-dependent selection gives an advantage to rare phenotypes, positive frequency-dependent selection gives an advantage to common phenotypes. In the between-species analogue, this is equivalent to an Allee effect, in which if a species is too rare, it may decline to extinction. This means that new alleles can have a difficult time invading a population, since they don't experience significant benefit until they become common.

Harmless scarlet kingsnake mimics the coral snake, but its pattern varies less where the coral snake is rare.

The effect can be seen in the evolution of warning coloration (aposematism) in toxic or distasteful organisms. The signalling theory advantage of such coloration is that predators can learn to avoid potential prey with that color pattern. For example, in the Batesian mimicry complex between a harmless mimic, the scarlet kingsnake (Lampropeltis elapsoides), and the model, the eastern coral snake (Micrurus fulvius), in locations where the model and mimic were in deep sympatry, the phenotype of the scarlet kingsnake was quite variable due to relaxed selection. But where the pattern was rare, the predator population was not 'educated', so the pattern brought no benefit. The scarlet kingsnake was much less variable on the allopatry/sympatry border of the model and mimic, likely due to increased selection since the eastern coral snake is rare, but present, on this border. Therefore, the color is only advantageous once it has become common.

See also

• Apostatic selection
• Evolutionary game theory
• Evolutionarily stable strategy
• Frequency-dependent foraging by pollinators
• Mimicry
• Tit for tat

Bibliography

• Robert H. Tamarin (2001) Principles of Genetics. 7th edition, McGraw-Hill.