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Kim Sneppen
The ¹elsBohr Institute, Blegdamsvej17, DK-2100 Copenhagen 8, Denmark
Per Bak
Brookhaven National Laboratory, Upton,
(Received 5 August 1993)
¹w York 11979
A simple dynamical model for Darwinian evolution on its slowest time scale is analyzed. Its mean
field theory is formulated and solved. A random neighbor version of the model is simulated, as is a
one-dimensional version. In one dimension, the dynamics can be described in terms of a "repetitious
random walker" and anomalous difFusion with exponent 0.4. In all cases the model self-organizes to
a robust critical attractor.
PACS numbers: 87. 10.+e, 05.40. +j
Introduction.— Life on Earth is presumably the most the first process is guided by a gradient, while the latter
complex dynamical system known; too complex for quan- is exponentially suppressed and only occurs because of
titative modeling, it seems. The usual strategy in that variance between individuals in a species. Viewed on its
situation is to focus either on a manageable subsystem or slowest time scale, then, evolution is discrete: species sit
special case or on just a few aspects deemed most impor- at local fitness maxima, and occationally a species jumps
tant for the entire system. The first approach was taken to another maximum. In the latter process, a species
in the outstanding work by Eigen and co-workers on pre- may change the fitness landscapes of other species which
biotic evolution [1]. We choose the latter approach here, depend on it, to the extent that some of them no longer
analyzing a quantitative model for Darwinian evolution find themselves at local maxima. Consequently, they im-
in general, with an eye on qualitative features of actual mediately jump to new maxima. This may affect yet
evolution. The model describes an ecosystem of interact- other species in a chain reaction, a burst of evolution-
ing species which evolve by mutation and natural selec- ary activity. We assume this chain reaction is subcritical
tion [2). Species spend most of the time at punctuated and on the average involves a total of K species (see [6]
equilibria [3,4]. Casually connected escapes of species for a realization of this situation), all in a time that is
from these equilibria form avalanches of evolutionary ac- negligible on the slowest time scale.
tivity with a power-law distribution of sizes. A mean field We characterize the state of an ecosystem of N species
theory yields an exponent ~ = 2 for this size distribution. by N values (x, ), i = 1, 2, . . . , N. These values charac-
The mean field description of the self-organized state is terize the effective barriers towards further evolution ex-
compared with a random neighbor version of the model perienced by the species at their local maxima of fitness.
and with a one-dimensional version. In one dimension, Since the waiting time for further evolution increases ex-
the dynamics can be described in terms of anomalous ponentially with the barrier height, the dynamics consists
diffusion with exponent 0.4. in selecting the species with the lowest barrier value, and
Simple as it is, the model has an ancestry: we have replacing that value, and that of K —1 other species, with
drawn on our earlier work on models of evolution inspired new values. For simplicity, we assume that the new bar-
by Kauffman's work [5—7], on self-organized criticality [8], rier values are random, all drawn from the same uniform
and on nonequilibrium growth of surfaces [9]. Also, we distribution in the interval [0, 1]. Results do not depend
are not the first to play with toy models of evolution; see, on this choice of distribution, as simple reparametriza-
for example, the last section of [10,11], and for a review tions relate all choices. Results do depend on the way the
[12]. K species are chosen. One choice consists in placing the
—
The model. We consider a dynamical ecosystem of in- species on the sites of a d-dimensional hypercubical lat-
teracting species which evolve by mutation and natural tice with nearest neighbor interactions. Thus K = 2d+ l.
selection. For simplicity, we assume that no species di- We shall return to the case of d = 1 below.
vide into several species and no species become extinct. Random neighbor and mean field model Here, for.
Thus the only effect of evolution is adaptation to the mathematical convenience, we select the K —1 interacting
environment. The environment of each species may be species at random among the N species in the ecology.
thought of as a fitness landscape with many local max- This random neighbor mode/ is a first step towards a solv-
ima, supplemented with some dependence on the states able mean field theory. We also assume this randomness
of some other species. We assume that evolution to lo- to be "annealed;" i.e. , the next time the same species
cal fitness maxima takes place much faster than escapes triggers K —1 other species to evolve, they are chosen at
from such maxima. This is a reasonble assumption, since random anew. A mean field theory can be constructed by
0031-9007/93/71 (24)/4087 (4) $06.00 4087
1993 The American Physical Society
VOLUME 71, NUMBER 24 PH YSICAL REVI EW LETTERS 13 DECEMBER 1993
p (x) =
1, N, P' (x)p(x)Q '(x) (1) o(*)=(' " ( -*)- ' "e*~)""
where we have introduced
= (1 —Kx) for 1/K —x )) O(1/N) .(9)
P(z) = dx'p(z'), Using p(x) = —&" Q(x), we have
K
=—
p(x) for 1/K —x » O(1/N), (10)
Q(x) = dx'p(x') .
distributed, p is left unchanged, as it should be. avalanches critical, our model for biological evolution is
This explanation points to another aspect of the a self-organized critical dynamical system.
asymptotic dynamics: If we trace in time which species One-dimensional model.— So far, we have seen criti-
trigger the bursts of evolutionary activity, then it is usu- cality only in the mean field approximation. Now let us
ally one of the species participating in recent activity. So study a finite dimensional case. We have simulated the
at any given late time, the very species which acquired dynamics of the one-dimensional ecology and measured
their current properties most recently are also the ones a number of its properties in the equilibrium state. Fig-
most apt to change them again. Thus, according to our ure 1(b) shows the distribution of barriers, p(x), and the
model, the cockroach, which is much older than the hu- distribution for the lowest barrier value, pi(x), as full
man race, will resemble itself long after humans, as we curves. Both are for K = 3, N = 100. They do not
know them, have disappeared. resemble the random neighbor and mean field results for
Avalanches. — In order to express the causal connec- K = 3, N = 100, shown in Fig. 1(a). The dashed lines in
tions between bursts of evolutionary activity, we define Fig. 1(b) show results from a difFerent mean field model,
an avalanche as a causally connected sequence of activ- obtained also with K = 3 and N = 100, but by replac-
ity associated with barrier values below the self-organized ing the two smallest barrier values plus one randomly
threshold 1/K. Suppose that at some time all barrier val- selected value with random numbers in each time step.
ues are above the threshold value. Then the next burst It is easy to understand why this latter algorithm gives
will, on the average, result in one barrier value below results much closer to the 1D results: Low barrier val-
threshold, which for its part will result in another barrier ues are clustered in one dimension, so the replacement of
value below threshold, etc. Thus the number of barriers the smallest barrier value together with the values on its
below threshold remains constant equal to one, on the nearest neighbor. sites amounts to replacing the lowest
average. The actual number of barriers below threshold value plus 0—2 other low values. Actually, some of the
fIuctuates and may become zero again, terminating the difI'erence between the mean field and 1D results shown
avalanche. in Fig. 1(b) is due to finite-N effects being more pro-
A more realistic value for the average number of barrier nounced in the 1D results, for example, the value of p(2:)
values below threshold can be obtained from our mean for x & 0.7. It will approach 3 as N — + oo, while the
field approximation. It gives mean field value for p(x) is already very close to 3 for
NP(1/K) = ln N —lnln N —ln(K —1) x) 0.7.
Figure 2 shows a space-time map of those sites on
+O(ln ln N/ ln N) which species change barrier values in the time inter-
+O(l/ ln N) + O(ln N/N), (12) val covered. Whenever the lowest barrier value is found
where P(1/K) = 1 —Q(1/K), and Q(l/K) is the so-
lution to Eq. (7) with x = 1/K. With an average of 200 T