What r and F actually mean
Coefficient of relatedness (r)
r is the probability that, at a random locus, two individuals share an allele *by descent* — that is, inherited from the same ancestral copy, not just the same sequence. Two completely unrelated mice might share a lot of allele sequences by chance (they are both mice!), but r counts only the shared-by-descent fraction. r = 0.5 means 50% of loci are identical by descent; r = 0.125 means 12.5% (first cousins).
Inbreeding coefficient (F)
F is the probability that, at a random locus, an individual is *homozygous by descent* — that both of its alleles trace back to the same ancestral copy in its pedigree. For the offspring of two parents with relatedness r, F_offspring = r / 2. That is why full-sib mating (r = 0.5) gives F = 0.25: a quarter of the offspring's genome is homozygous by descent.
Why it matters in the colony
Inbreeding depression — reduced litter size, smaller pups, lower fertility — starts to become detectable around F ≈ 0.0625 in outbred stocks and gets progressively worse as F climbs. The NIH Guide for the Care and Use of Laboratory Animals treats the first-cousin threshold (F = 0.0625) as the practical boundary. Maintained inbred strains operate well above this by design and use the homozygosity as the whole point of the strain.
Wright's path coefficient method
The formula
For a pairwise relatedness between X and Y traced through a single common ancestor A:
r(X, Y) = (0.5)^L × (1 + f_A)
where L is the number of links (generation steps) in the path X → A → Y, and f_A is the inbreeding coefficient of the ancestor itself.
Walking the path
Full sibs share *two* common ancestors — both the dam and the sire. Each path is length 2 (X ← parent → Y), contributing (0.5)^2 = 0.25. Two paths sum to 0.5.
First cousins share one pair of grandparents. Each path is length 4 (X ← parent ← grandparent → parent → Y), contributing (0.5)^4 = 0.0625. Two paths (one through each grandparent) sum to 0.125.
The (1 + f_A) correction
If the common ancestor is itself inbred, the two alleles passed down the two sides of the pedigree are more likely to already be identical. This inflates r by a factor of (1 + f_A). For non-inbred founders, f_A = 0 and the correction vanishes — the standard textbook values assume this.