SLIDE 8
- Inheritance to the next generation: If we know relatedness in one generation,
we can compute relatedness in the next generation. Suppose x and y mate to produce z. Then for any w other than z itself or a descendant of z, F(w, z) = 1
2F(w, x) + 1 2F(w, y).11
Why? Consider a random allele g from w (if you like, one of w’s two body
- dor alleles). How many copies does z have on average? Well, the mother
x has F(w, x) copies on average, of which z inherits half on average.12 And the father y has F(w, y) copies on average, of which z also inherits half on average. As an example of particular interest, consider w = x, which tells us how much x shares with her own child z: F(x, z) = 1
2F(x, x)+ 1 2F(x, y). Without
inbreeding, F(x, x) = 1 and F(x, y) = 0, so this works out to 1
2, as assumed
in question 2b. But the formula rightly expects it to show up more often (i.e., F(x, z) > 1
2) if an allele g from x is likely to appear more than once in
x (i.e., F(x, x) > 1), or is also likely to appear in y (i.e., F(x, y) > 0). Note that this example requires us to know F(x, x), which measures how
11As a sanity check, note that if F(w, x) and F(w, y) are both in the range [0, 2] as they’re supposed to
be, then so is F(w, z). As another sanity check, suppose w’s parents are u and v. Then there are two ways to compute F(w, z) (provided that all “no descendant” conditions are met), and fortunately they give equal results: F(w, z) = 1 2F(w, x) + 1 2F(w, y) = 1 2(1 2F(u, x) + 1 2F(v, x)) + 1 2(1 2F(u, y) + 1 2F(v, y)) F(w, z) = 1 2F(u, z) + 1 2F(v, z) = 1 2(1 2F(u, x) + 1 2F(u, y)) + 1 2(1 2F(v, x) + 1 2F(v, y))
12In other words, z inherits each of x’s copies of g with probability 1
- 2. We are assuming that the fact that
g is known to appear in w does not make z any more or less likely to inherit that gene. This assumption fails in the case where w happens to be z, in which case z inherits g with probability 1. That is why w is not allowed to be z; we will handle the case w = z separately. More generally, the assumption fails if w is any descendant of z, even if w is not z itself. In that case, when we are trying to guess whether z inherited from x a given one of x’s copies of g, the probability is > 1
2
if we know that g also made it down to z’s descendant w. The assumption fails in the same way if w happens to be z’s identical twin, in which case z again inherits g with probability 1, or a descendant of z’s identical twin. The easiest way to handle identical twins is to pretend they are the same person, i.e., give them a single node in the family DAG, though this node may have roughly twice the typical number of mates and offspring. Biologically speaking, are we otherwise on safe ground? Almost. It is true as far as I know that inheritance events are independent—i.e., if you have allele g, you really are 1
2 likely to pass it on to each of your gametes,
independent of who you inherited it from or which of your relatives got it too. However, it may be that you are not 1
2 likely to pass it on to each of your children, since perhaps g is maladaptive (either by itself or in
combination with certain other alleles of the same gene or other genes) in such a way that the gametes with g are less likely to lead to actual births. Thus a child z that actually shows up in the family DAG as a birth may have < 1
2 chance of having g (and > 1 2 chance of having your other allele). We ignore this “selection
effect.”
8