The Hardy–Weinberg principle, also known as the
Hardy–Weinberg equilibrium, model, theorem, or
law, states that allele and genotype frequencies
in a population will remain constant from generation to generation
in the absence of other evolutionary influences.
The seven assumptions underlying Hardy–Weinberg equilibrium are
as follows:
- organisms are diploid
- only sexual reproduction occurs
- generations are nonoverlapping
- mating is random
- population size is infinitely large
- allele frequencies are equal in the sexes
- there is no migration, mutation or selection
Violations of the Hardy–Weinberg assumptions can cause
deviations from expectation. How this affects the population
depends on the assumptions that are violated.
- Random mating. The HWP states the population will have the
given genotypic frequencies (called Hardy–Weinberg proportions)
after a single generation of random mating within the population.
When the random mating assumption is violated, the population will
not have Hardy–Weinberg proportions. A common cause of non-random
mating is inbreeding, which causes an increase in homozygosity for
all genes.
If a population violates one of the following four assumptions,
the population may continue to have Hardy–Weinberg proportions each
generation, but the allele frequencies will change over time.
- Selection, in general, causes allele frequencies to change,
often quite rapidly. While directional selection eventually leads
to the loss of all alleles except the favored one (unless one
allele is dominant, in which case recessive alleles can survive at
low frequencies), some forms of selection, such as balancing
selection, lead to equilibrium without loss of alleles.
- Mutation will have a very subtle effect on allele frequencies.
Mutation rates are of the order 10−4 to 10−8,
and the change in allele frequency will be, at most, the same
order. Recurrent mutation will maintain alleles in the population,
even if there is strong selection against them.
- Migration genetically links two or more populations together.
In general, allele frequencies will become more homogeneous among
the populations. Some models for migration inherently include
nonrandom mating (Wahlund effect, for example). For those models,
the Hardy–Weinberg proportions will normally not be valid.
- Small population size can cause a random change in allele
frequencies. This is due to a sampling effect, and is called
genetic drift. Sampling effects are most important when the allele
is present in a small number of copies.