Chapter 20
The diffusion of genes (1937)
In 1937, two articles were published introducing a new approach to the study of spa-
tial heterogeneity in population dynamics. Fisher was the author of the first article,
entitled The wave of advance of advantageous genes, which appeared in the Annals
of Eugenics. He studied the spatial propagation of a favourable gene in a popula-
tion. As a simplification, he considered a space reduced to just one dimension and
called u(x,t) the proportion of the population located at point x at time t that pos-
sesses the favourable gene. So 0 ≤ u(x,t) ≤ 1. To include natural selection, he used
equation (14.6) with a continuous time variable
∂u
= a u (1 − u) ,
∂t
where a is a positive parameter. For a given value of x, we recognize Verhulst’s
logistic equation (see Chapter 6) with a solution u(x,t) that tends to 1 as t → +∞.
Furthermore, Fisher assumed that the offspring of an individual located at point x
with the favourable gene do not stay at the same point but disperse randomly in
the neighbourhood of x. By analogy with physics, he argued that one must add a
diffusion term to the equation for u(x,t), leading to the partial differential equation
∂u ∂ 2u
= au 1−u +D 2 . (20.1)
∂t ∂x
When the selection coefficient a is zero, this reduces to the diffusion equation in-
troduced by Fourier in his theory of heat and later used by Fick for the diffusion
of physical particles. In 1904, Ronald Ross had started considering random disper-
sal in population dynamics. He was then wondering how the density of mosquitoes
decreases as the distance from a breeding site increases. The problem had come to
the attention of Karl Pearson and Lord Rayleigh. By 1937 the body of scientific
literature on diffusion equations had grown considerably, in particular following
Einstein’s work on Brownian motion.
Fisher showed that there exist solutions of equation (20.1) of the form u(x,t) =
U(x + vt) satisfying the three conditions
N. Bacaër, A Short History of Mathematical Population Dynamics, 111
DOI 10.1007/978-0-85729-115-8 20, © Springer-Verlag London Limited 2011
112 20 The diffusion of genes (1937)
0 ≤ u(x,t) ≤ 1, u(x,t) −→ 0, u(x,t) −→ 1,
x→−∞ x→+∞
provided that v ≥ v∗ where √
v∗ = 2 a D .
These solutions connect the steady state u = 1 with the favourable gene to the steady
state u = 0 with no such gene. They represent waves propagating at speed v in the
direction of decreasing values of x. Indeed, u(x − v T,t + T ) = u(x,t): the part of the
wave that was in position x at time t moves to position x − v T at time t + T .
Fig. 20.1 Propagation from
left to right of a favourable
gene at the speed v∗ . The gene
frequency u(t, x) at t = 0 is a
step function.
∂u
Indeed, setting z = x + vt, Fisher noticed that if u(x,t) = U(z), then ∂t =
vU (z), ∂∂ ux = U (z) and ∂ 2u
∂ x2
= U (z). If u is a solution of equation (20.1), then
vU (z) = aU(z) (1 −U(z)) + DU (z) . (20.2)
When u is close to 0, i.e., when z → −∞, Fisher expected that U(z) → 0 and
U (z) → 0. Calling k the limit of U (z)/U(z) when z → −∞, we know from
L’Hôpital’s rule that U (z)/U (z) also tends to k. Therefore, U (z)/U(z) =
[U (z)/U (z)]×[U (z)/U(z)] tends to k2 . Dividing equation (20.2) by U(z) and
letting z tend to −∞, we arrive at a second-order equation D k2 − v k + a = 0.
But k must be a real number. So the√discriminant of this equation has to be
nonnegative: v2 − 4 a D ≥ 0, or v ≥ 2 a D = v∗ . Hence, v ≥ v∗ is a necessary
condition for the existence of a wave propagating at the speed v. It is also a
sufficient condition, as explained below.
Fisher noticed that only the wave which propagates exactly at the speed v∗ is
selected for a large class of initial conditions, e.g., for the step function: u(x, 0) = 0
for x < 0, u(x, 0) = 1 for x ≥ 0. Figure 20.1 shows how this discontinuous initial
