Inbreeding and Outbreeding

Contemplate the descent of a piece of DNA (or RNA in organisms using this as their genetic material). The DNA is copied, and copies are passed to descendants. If the copies were error-free we could rightly think of them as perfect clones that pass down indefinitely through the eons. This logic led Richard Dawkins to speak of immortal coils in his book on selfish genes; here, it instead brings up issues of the common ancestry of genes and of individuals, and of the definition and consequences of inbreeding and outbreeding, the subjects of this chapter. When two individuals share one or more ancestor, they are relatives, both in common parlance and by technical definition in biology. The consequence of their mating is inbreeding, that is, the production of offspring receiving copies of a given gene through both mother and father that can be traced to the common ancestor(s). These gene copies are identical by descent (IBD; not to be confused with an acronym for inbreeding depression, see below), a shorthand for “identical by the fact of descending as copies of the same original piece of DNA”. The probability that two gene copies are IBD in a diploid individual, or its inbreeding coefficient, symbolized by f, is a simple function of the genetic relatedness of its parents and the segregation of genes during meiosis and gametogenesis. Because the probability is one-half that two gametes from the same individual carry identical gene copies, fertilization by self produces f of one-half, a brother-sister mating or parent-offspring mating produces f of one-quarter, a first-cousin mating produces f of one-sixteenth, and so on (see “Measurement of Inbreeding and Outbreeding,” below). In these examples, we assume that neither common ancestor(s) nor parents themselves are inbred; such inbreeding reflects additional common ancestry and so inflates f. From all of this, a definition of outbreeding as “mating of nonrelatives” follows automatically. As just defined, inbreeding and outbreeding rely on an absolute measure of relatedness. An alternative definition that may be of more value in real, finite populations (as opposed to ideal, infinite ones) is that inbreeding is mating with relatives more often than expected by chance, and outbreeding the opposite.