What are inbreeding and linebreeding, and what effect do they have?

In genetic terminology, inbreeding is the breeding of two animals who are related to each other. In its opposite, outcrossing, the two parents are totally unrelated. Since all pure breeds of animal trace back to a relatively limited number of foundation dogs, all pure breeding is by this definition inbreeding, although the term is not generally used to refer to matings where a common ancestor does not occur behind sire and dam in a four or five generation pedigree.

Breeders of purebred livestock have introduced a term, linebreeding, to cover the milder forms of inbreeding. Exactly what the difference is between linebreeding and inbreeding tends to be defined differently for each species and often for each breed within the species. On this definition, inbreeding at its most restrictive applies to what would be considered unquestioned incest in human beings - parent to offspring or a mating between full siblings. Uncle-niece, aunt-nephew, half sibling matings, and first cousin matings are called inbreeding by some people and linebreeding by others.

What does inbreeding (in the genetic sense) do? Basically, it increase the probability that the two copies of any given gene will be identical and derived from the same ancestor. Technically, the animal is homozygous for that gene. The heterozygous animal has some differences in the two copies of the gene Remember that each animal (or plant, for that matter) has two copies of any given gene (two alleles at each locus, if you want to get technical), one derived from the father and one from the mother. If the father and mother are related, there is a chance that the two genes in the offspring are both identical copies contributed by the common ancestor. This is neither good nor bad in itself. Consider, for instance, the gene for PRA (progressive retinal atrophy), which causes progressive blindness. Carriers have normal vision, but if one is mated to another carrier, one in four of the puppies will have PRA and go blind. Inbreeding will increase both the number of affected dogs (bad) and the number of genetically normal dogs (good) at the expense of carriers. Inbreeding can thus bring these undesirable recessive genes to the surface, where they can be removed from the breeding pool.

Unfortunately, we cannot breed animals based on a single gene - the genes come as a package. We may inbreed and rigorously remove pups with PRA or even their parents and littermates from the breeding pool. But remember inbreeding tends to make all genes more homozygous. In at least one breed, an effort to remove the PRA-causing gene resulted in the surfacing of a completely different and previously unsuspected health problem. It is easier and faster to lose genes (sometimes very desirable genes) from the breeding pool when inbreeding is practiced than when a more open breeding system is used. In other words, inbreeding will tend to produce more nearly homozygous animals, but generally some of the homozygous pairs will be "good" and others will be "bad".

Furthermore, there may be genes where heterozygosity is an advantage. There are several variant hemoglobin types in human beings, for instance, where one homozygote suffers from some type of illness, the other homozygote is vulnerable to malaria, and the heterozygote is generally malaria-resistant with little or no negative health impacts from a single copy of the non-standard hemoglobin gene. A more widespread case is the so-called major histocompatibility complex (MHC), a group of genes where heterozygosity seems to improve disease resistance.

Is there a way of measuring inbreeding? Wright developed what is called the inbreeding coefficient. This is related to the probability that both copies of any given gene are derived from the same ancestor. A cold outcross (in dogs, probably a first-generation cross between two purebreds of different, unrelated breeds would be the best approximation) would have an inbreeding coefficient of 0. Note that this dog would not be heterozygous at every locus. There are genes shared with every multicellular organism, genes shared with all animals, genes shared with all animals with backbones, genes shared with all four-limbed animals (including most fish and all amphibians, reptiles, birds and mammals) and with all mammals. Although the DNA might differ slightly, the proteins produced would be functionally the same. Further, the chances are that our dogs with inbreeding coefficient = 0 would still be homozygous for some genes shared by all dogs. The inbreeding coefficient thus specifically refers to those genes that are variable (more than one possible form) in the species and even the breed being considered.

An inbreeding coefficient of 1 (rare in mammals) would result if the only matings practiced over many generations were between full brother and full sister.