Subscribe to DeepL Pro to translate larg

Visit www.DeepL.com/pro for more in

J.Z. Fisher's Equation and solutions of the propagating volim type 213

here, they are still, in essence, kshematic waves, since their propagation is
caused by pelinear ones
"reaction" terms in the right-hand side of (5.10).

5.3. Uriaviete Fvuera

n propagating wave grids
The main equation studied here i s an hour example of a class of scalar
reaaa tion equations with diffusion (5.1 in one-dimensional pro- stranse
and, ku l u) -1- Dng , (5.12)
where the saaalar function p (x, d) satisfies the given initial and boundary
conditions, and k z D is a constant. This equation is important from the
historical and pedagogical points of view. It was proposed (as well as the
equation with cubic nonlinearity instead of avadshchtic in the right part) by
Fisher (1937) as a deterministic version of the spchastic model of the spread of
a favourable gene in a diploid population. He examined the equation in detail
and obtained a number of useful results, the derivation and application of which
we will demonstrate below. A. N. Kolmogorov, I. G. Petrov-siai, and N. S.
Piskunov, whose classic work (1937) served as the basis for a more cmpororo
analytical approach to the Fisher equation, also gave a heuristic and re- netic-
based derivation of the equation. In recent years, Aronson and Weinberger
(1975), McKeon (1975), Fyfe and MacLeod (1975, 1977), and Larson (1977)1 '
have considered a broader class of equations in which the yalei ku(1 and) is
replaced by a szalvryai function F in) belonging to a non-natural class of
functions. The Fisher equation is one of the simplest nonlinear equations of
reactions with diffusion, in which at least one of the types of waves of interest
to us, namely, the damped front, arises, so it is so important pedagogically
and therefore it is given here so much space.
Equation (5.12) is the simplest diffusion model for the logistic model of the
population growth, which is touched in the prior partition R, which gives
(undiffusive) relations o f t h e kinematic wave type. Therefore, the main
purpose of the study of equation (5.12)-determine the influence of diffusion on
the ktematically propagating waves observed in the absence of diffusion. The
wave solutions of Eq. (5.12) are essentially still kinematical. We want to
investigate the existence and form of solutions of the travelling-wave type of
e q u a t i o n (5.12), for which 0 p< z 12 ', and find the
velocity

' See also Stokes (1976)*, Rothe ( 1978) *, and yanggu

(1979) *.- P rim. transl.
" In Fisher's (1937) equations probability.
214 Chapter 5. Biological Oscillators

the propagation of such a war. If a solution to the running warfare exists, it can
be written in the form of

where c is the aolne velocity. Since equation (5.12) is supported by replacing x

by x, the ratio c can be positive or negative; for definiteness, we will assume
c to be positive, so that (5.13) represents a wave travelling in the negative
direction of the x-axis. After fitting (5.13) to (5.12), the function satisfies
the equation
Df" cf' + kf[1 0, (5.14)
where shtryah means differentiation over Z. Since (5.12) is invariant under
constant displacement in x and t, an arbitrary constant can be added to z in
(5.13). We now want to find an eigenvalue or values of c such that equation
(5.14) has a non-perturbative solution for which
lim - 0, lish 1.

An important additional question is as follows: if such a solution exists,

then for any initial profile for which 0 -3 Ts(x, 0) fi 1 at each x, the
corresponding solution of Eq.
{5 12) satisfying the boundary conditions in t) 0, tsr ) - 1,
as t grows, the travelling wave type solution (5.14) satisfying the
that fulfils the conditions (5.15)? .
Fmep (1937) found that equation (5.14) has an infinite number of
travelling wave type solutions, for which 0 in y 1, with the following
values
с c"" - 2 (5.16)
We will show it below. A.N. Kolmogorov, I.G. Petrovsky, and N.S. Pi-
(1937) have reported that the lry

1, xc < x,
х < 0,
m(x, 0) andmi and;x, 0) y(x), x2 < x < xJ , (5.17)
х > 0,
0, х < 2 '
where xi and x2 are coichnv, and h(x) is monotone and continuous, (5.12) has
an edt-whence solution and this solution evolves into a solution {5.14) of the
monotone running wave form satisfying (5.15) and having a speed with cp,п
. A typical herein solution is shown in Fig. 5.3.
Generally speaking, the behaviour and{x,0) at x + is decisive
8

for the evolution of 6erymx waves in time. This question is specifically

addressed to-
 5.3. Fisher's Uraanenw# Fisher and propagating type solutions herei 215

The problem has been addressed by Maakin (1975\ Larson (1977) and Fyfe
and MacLeod (1977). We will point out the difficulties that arise in this case
when we discuss the stability of t h e travelling wave type solution.
To show that for c and c"" there exist wave solutions in the form (5.13), let
us use the phase plane method for the equation (5.14) (see, for example, the
books by Minoschky ( 1962) or Sansone and Conti (1964)) and look for the
conditions under which there exists a noncontinuous solution satisfying the
conditions 0 1 and the
boundary conditions (5.l-5). Thus, for c we obtain the problem on the
eigenvalue in an infinite domain.
Ma can write {5.14) in the form
f F, DF - cF k/(1 f),
then the trajectories in the phase plane (/ F will be solutions ypame-

dF cF k/(1
(5.18}
df DF

which has two special points in the plane: (0, 0) and (1, 0). The wave solution of
equation (5.12) of the type shown in Fig. 5.3 corresponds to the trajectory from
(0s0) to (1, 0) remaining in the band 0 ';; 1 with derivative F y 0.
Near the point (0, 0) equation {5.18) can be linearised, which gives

dFcF
kf DF
opuda (0, 0) represents co6oii iestvayu y zed if and only if and only if .shli s y
sudu 2 kD , i.e. (5.16). Esuii 0 k s c cq, special point ( 0, 0)
is an unstable focus, while at c -- 0 it is the centre; in these cases, the solutions
of f satisfying the required conditions are not
can be, so Eac there will be pointsR in any trajectory at (0, 0),
on which < 0.
Another non-conditional condition is that the singular point
(1, 0) is saddle-shaped. Near (1, 0) zh 1, and the lteaschised form (5.18) is here

dF cF -1- k I
DY
which corresponds to the saddle point for acex with y 0, since k and D are
polar constants. The trajectories in the phase plane are shown in Fig. 5.4. We
assume that for each yYi, 2 JYЇi there is a single trajectory starting at the
point 0 and moving towards the centre of the phase plane.
- 1 in the band 0 4 y 1, at which F > 0, except for points - 0
316 Chapter 5. Biological oscillatorsi

1, where N 0. At c rp, the knot at point (0, 0) is degenerate with two

slopes with - 1 - (c 2 4#D)1 '2 ]/2D, slvaying /2D - / .
A.N. Kolmogorov, I.G. Petrovsky, and N.S. Piskunov (1937) proved
nontrivially that cp,п is the natural speed of wave propagation, and the solution
of the Fisher equation (5.12) is obtained if the initial data have the value
(5.17). By "naturalness" we mean that the solution with such initial data will
evolve into a travelling wave, which is naturally shown in Fig. 5.3, which h a s
the same behaviour as rp; 2 [YЇi. Kaa mentioned above, the behaviour of
the solution over a large interval

Fig. 5.4. Wave trajectories

of new reactions of equation
(5.14) in phases and planes

}c 1 (c2 - 4kD)'')l2D.

The time depends crucially on the behaviour of the initial data

at infinity.
A simple heuristic reasoning that illustrates that the i n i t i a l data, expressed
in fact by the first of the initial conditions (5.17), will evolve into a wave solution
with with in - belongs to Fischer (1937). The idea, which has a wider
applicability, i s that the solution and x t) is first of all assumed to evolve into a
wave solution of constant schematically shown in Fig. 5.3. At any time d
the total
the area under the curve to the left of the point x - R 0 is equal to
-R
U and(x, t) dx,R > 0. (5.19)

In the reaction scheme with diffusion, U gives the total amount of reactant ts to
the left of x - R. finite t and sufficiently large I, the values in the left of R
will be small; for the first variant of the initial data (5.17), n is initially even zero.
The idea of Fischer (1937) was to fix some arbitrarily small value of U and to
define R as a function of time so that U remained equal to this prescribed value.
Thus, when U is fixed, the function R (t) is determined from (5.19) and, hence, can
be defined from (5.19).
 5.3 Fisher's Fisher's Udovnechts# and the solutions of the propagationR 8oyans 217

the propagation velocity dR(r)/dl can be found. For small u and large A mm we
are in the region where 0 < and 'to 1, so2 can be neglected in comparison with c,
and equation (5.12) is approximated by the linear equation

and, ku -I- Du . (5.20)

By solving this equation for the first variant of the initial conditions (5.17) and
estimating A(f) or the bestSchS dR/dt from (5.19} we can determine the
velocity of VOLI.
However, the maths is much simplified if we follow what Fischer (1937)
actually did. He considered a two-dimensional symmetric problem with g 0 as
a radial coor- dinate and the fun- tion ts and{g, t$ satisfying instead of (5.20)
Eq.

ii - ku -1- D up -1- and , p(r, 0) - fi (Jj, (5.2\)

where 6(¿)is the usual Dirac delta function ". Now instead of (5.19) we require
that the area under and outside the circle r L is constant, i.e.

2n tp (d, t) rdr - U - const. (5.22)

- = R (t\

To solve (5.21), (5.22), we write

this problem has a fundameital solution (see, e.g., Ky- rant and Hilbert ( 1962))
1
n(d, t)
4nDi
Thus, fl (t) can be obtained by R3 (5.22) after substituting
in from the last equation, i.e., as a solution of Eq.

') This is not a fusion in the general sense of the word. If /(x)-function from x, not
discontinuous at z - 0, fi (z) is defined by J f i (x)f(x)f(x)dx --f(0) for
zyubozo s n 0. Euristics 6(x) is zero, except for the race x = 0, rne it is unco
is finitely Velnha, and the last integral makes sense. Similarly reduced
218 Chapter 5. Biological Oscillators

it's after the integration

Ya(t) - 4D (ln U) t -- 2 t yu JÏ(d) - dYa/dl 2

Consequently, the speed of will spreading is set equal to -
|/ÈÈ', with
Л
, from (S.16).
Instead of taking (5.22) as a definition of R [t), we can define the position R (t)
of the wave as the position where u takes a fixed value Ïi, not necessarily small;
the natural choice is m '/ . Then R(/) is found from Eq.
- 1
*" 4nDt
instead
R2 (E)-- 4kDt' 4Dr in 4xyDr.
The propagation rate dR/dl for large times is found by diffe- renting:
dI
2 +О greater than t), (5.23)
dt
which indicates the order of the asymptotic correction to the wave velocity.
The analogous analysis applied to (5.19) and (5.20), with initial data
corresponding to the first variant of the conditions (5.17), requires the
integrals of the standard asymptotic methods ( see Eqs,
See, e.g., Murray ( 1974)). Again for boliysh | x | and t we obtain

'"' (') 2 О
а' "
In the question of the existence of wave solutions with velocities with >
*>in meogokrano has been discussed since the possibility of this was
recognised by Fischer (1937) and A.N. Kolmogorov et al. (1937). In recent
years, interest in the asymptotic study of travelling-wave type shocks and their
stability has increased considerably; we have given references to the relevant
works above. The paper by Mollison (1977), among others, discusses the
Fisher deterministic model in connection with the spatial contact model; this
model was considered only as an approximation of the more realistic stochastic
model. Although Molleason's (1977) paper is devoted more to stochastic
phenomena, he makes some interesting judgements about deterministic
diffusioinms
models and their relation to stochastic ones. He shows sufficient-
but in a simple way, as the war rate of the established solution

1' This result is also obtained under the second variant of the conditions (5.17), even

if £(x) is not required to be monotonic and uninterrupted.

 5.3 The Fiu#ry equation for a travelling wave solution 219

of the travelling wave depends on the initial conditions; we will give here
the main content of theero analysis.
As has seen, the assumption of Fäschsr (1937) that the wave velocity
depends on the leading edge of the wave, where in is small, was used to determine
this velocity. Once again, we assume u2 to be negligibly small and start with the
stump equation (5.Y). Consider the right-hand side of the equation

and(x,0) e°", and(x, t) e°("" "'', (5.24)

The second of which represents the solution in the form of a wave travelling to
the left with a velocity of c, Let us note the front R or the leading edge of the wave
as the zone in which ts is small, i.e., x + c' < 0 and } x + c' | vetko. Substituting
the variation for the travelling wave form (5.24) into equation (5.20) and
subtracting the exponential, we obtain the dispersion relation with k -1- Da2 for
the still
dependence of c[a), with the minimum velocity is equal
to
"s - 2 ; this is the velocity at which L means p from the diYazone 0 < o < up
voshovaya velocity c > s "i,.
Let us now consider miN {e°", e "*} at x to 0 and note that, for the linear
equation (5.20), they depend monotonically on the real conditions. If o < su,
then for x < 0 e°" > e°°', so that the propagation velocity with the initial condition
(5.24) and with such p will depend on the front or leading edge of the wave. On
the other hand, the initial conditions C, d o0 require that e°' be bounded from above
by the function e'°' at x < 0, and so the propagation velocity will depend on the tail
of the wave. In other words, if f o r the Fisher equation (5.12) and{x,01 S
4 e"' at x -+ 'y for some o п lx, d) enclosing
but by a pseudowave with speed sudu, namely e'°"+ kfD, '"" for all i "
" 0 ' . ShR conclusions apply to the case of more than one
of the spatial dimension.
The results presented here can be generalised to a class of equations-
iejayi react"-* - n-b-8 zyed species
and, F{i) + Du, , (5.2S)
where n is a link and F{i) is continuous for 0 fi and 1 и

F {i) du > 0, F(0) - - - f (l) - 0, F {1} 0. (5.26)

Esh equations have been discussed in detail by Aronsoy and Weinberger (1975),
Huds-.
In the case of the study, it was used in the traditional methods of phase

1 This reasoning remained a mystery to the writer. However, we decided to

give their exact translation in the hope that some of the readers will be more clever than
others. The prize.
 220

The latter model, Fife and MacLeod (1977) and Larson (1977), contains
references to epidemic waves and early generalisations of the Fische equation,
especially for some combustion problems and to models of nerve impulse
propagation in which travelling wave solutions also arise. In the latter model F
'and) and I p) (h u), where 0 < a < 1. (5.25), (5.26) with the help of a
simple development of the analysis demonstrated above with the use of the
phase plane method, it can be shown that wave solutions have a meaning and
exist only if c > co;. 2 DF 0).
KaE mentioned above, since the Fisher equation (5.12) inva-
is variable with respect to the change of the sign of x, there is a solution of the
wave type,
running and right: ts(x, t) - /(x ct\c > 0, where now 1и

Rns. 5.5. Time evolution of a typical solution of the Fisher equation (5.12), which ensures
the constancy of the splanchnic fronts.

0. It is therefore natural that if we start with a finite polar perturbation in

which p {x, 0) 0 outside the finite region, the waves will move in both directions, as
shown in Fig. 5.5.. We notice that if for those x where p (x, 0) > 0, p (x, 0) < 1 is
fulfilled, then the term ku!l u) leads to an increase in the p e r t u r b a t i o n ,
so that lim and[x, d) - l for the
of all x. We can simply consider Ok (1 n) as a LOGICAL STOCHRICK
in the diffusion equation when c0 , however small it may b e .

On the role of the Fnscher equation and the work of Fi. N. Kolmogorov, I. G.
Petrovsko-go and N. S. Piskunov (1937) on the role of the Fnsher equation and the work of
fi. N. Kolmogorov, I. G . Petrovsko-go and N. S. Piskunov (1937).M. Mihvaladze (1980)*
(mathematical results are also given in the articles by I. M. Gepfatzda (1959)* and G. I.
Barenblatt and J. B. Zeldovpyaa (1971)*). transl.