Genotype fitness 2026-10-13
Genotype fitness measures the reproductive or viability success assigned to a genotype in a specified selection model. Relative fitnesses weight Hardy-Weinberg proportions after viability selection; continuous-time fitness differences instead act as selection-rate differences. The convention and time units must be specified before comparing selection with migration.
Consider an explicitly idealized haploid population with two inherited types , one allele per individual at the tracked genetic locus, and discrete, nonoverlapping generations. Assume faithful type transmission, no mutation or gene flow, a constant environment, and reproductive contributions independent of type frequency. Let the expected values of offspring surviving to the next census be , with relative genotype fitness .
For frequency , expected offspring contributions are proportional to and . Normalizing gives the deterministic large-population mathematical model
The denominator is mean relative genotype fitness. For and , . Equivalently, 200 and 800 parents with contribute expected counts 240 and 800. Fractional per-parent expected values are averages, not literal fractional offspring. The ratio of these expectations is a deterministic approximation, not generally the expectation of a random ratio.
The odds multiply particularly simply:
Thus : repeated modest advantage can substantially change composition. For and , . The deterministic boundary remains unchanged: natural selection needs existing variation.
Figure 1.
Deterministic selection and stochastic sampling in a finite population
. Both panels start at . The left compares the deterministic haploid selection recurrence with eight simulated finite-population paths at ; the right shows eight neutral genetic drift paths at . Each simulation samples offspring independently per generation. These are illustrative mathematical models, not experimental measurements. Details.
A distinct stochastic extension fixes offspring and samples their types independently using a binomial distribution: , where , then . Its conditional expected value is and conditional variance is . With , the conditional expected value remains , yet realized frequencies wander through genetic drift. With , genetic drift can still lower or lose entirely. Over several generations, the deterministic curve need not equal the mean stochastic trajectory, because the recurrence is nonlinear.
Heterozygote advantage 2026-10-13
A heterozygote has greater genotype fitness than either corresponding homozygote. In the deterministic random-mating two-allele viability-selection model this gives an attracting interior allele frequency, unlike the repelling interior frequency produced by underdominance.
Natural selection 2026-10-13
Natural selection involves systematic differences in expected reproductive success associated with variation in phenotype. For a continuing evolutionary response, the relevant variation must be inherited: offspring must tend to preserve the differences being favored. Actual offspring counts also fluctuate by chance, producing genetic drift. Survival matters insofar as it affects contribution to future generations. Genotype fitness is therefore contextual, depending on environment and the life cycle being counted, rather than simply measuring strength or longevity. See Clark, Douglas and Choi, Biology 2e: population evolution.
In chapter IV of his 1859 book, Darwin explained natural selection through preservation of favorable variations and greater chances of survival and reproduction. See Darwin, On the Origin of Species, first edition, chapter IV. The beautiful consequence is cumulative change without foresight: inherited differences are filtered repeatedly by present conditions. Natural selection neither generates a needed mutation on demand nor guarantees progress toward a predetermined goal.
Underdominance 2026-10-13
Underdominance means that the heterozygote has lower genotype fitness than both corresponding homozygotes. In a deterministic randomly mating biallelic population, it produces an unstable interior allele-frequency threshold separating the two fixation outcomes, unlike heterozygote advantage.
With homozygote fitness advantages over the heterozygote, the allele frequency of the first allele has an unstable threshold . Below it that allele decreases; above it it increases. In a discrete viability-selection generation the same numerator gives after division by mean genotype fitness. Equal homozygote advantages give , with stable endpoints and unstable midpoint.