A model of evolution

M. Bunge
Foundations and Philosophy of Science Unit, McGill University, Montreal H3A1W7,
Canada
(Received March 1978; revised May 1978)

The theory of evolution is too general to be able to make predictions.

However, like every other hypergeneral scientific theory, it can be
enriched with specific assumptions and data, and thus become a
specific model for a particular evolutionary' line. This paper exhibits
such a model. It accounts for the evolution of a biopopulation in
terms of genetic and environmental variables. The result is a system
of linear finite difference equations that enable one to pin down the
necessary and sufficient conditions for evolution proper to occur, as
well as to predict the average numbers of the final varieties
originating from the parent population, it is estimated that the main
value of the paper is methodological rather than substantive, in that it
suggests a way of building models of biological lineages.

The synthetic theory of evolution is usually said to be Phenomenological model

incapable of formulating predictions, hence of being
tested, ergo of qualifying as a scientific theory. There is
a grain of truth in this overstatement; that, unless that
theory is adjoined to a precise model of the organisms
that are of interest, it is so general that nothing can
be predicted with it. The theory is very general but it
can and must be applied to specific lineages if we wish
to make evolutionary predictions.
At first sight nothing could be easier than setting up
specific evolutionary models. However, the dearth and
one-sidedness of such models in the literature suggests
that things are not easy. In effect the main difficulty
seems to lie in the choice of components of the state
function that spans the state space of a population (or
ecosystem) as the latter evolves. There are three
possible choices. One may choose the gene frequencies
as the coordinates Or components of the state function.
This is the course adopted by population geneticists. It
has not been over fruitful because selection operates on
phenotypes or molar variables rather than genotypes.
Another obvious choice is to focus on molar or
phenotypic variables. But this would leave the primary
source of individual variability in the dark. And neither
of the above strategies allow for environmental
variables.
The way out of this impasse seems to be to take
genetic as well as environmental variables into account.
More precisely, we suggest taking the census counts of
the various subpopulations of a given biopopulation in
the course of time; and viewing them as the net effects
of genic variability and environmental selection, We
will show, in the idealized example below, how this
program can be implemented.
We propose to sketch a simple model of the
evolutionary process. Consider a biopopulation of
organisms, such as bacteria of some kind, that
reproduce by binary division. Assume that each
individual lives one time unit, at the end of which it
divides simultaneously with all the other members of
the population. Assume also that mutants appear at
each generation and that they can be grouped into
distinct varieties. Call V~the ith variety, where
1 ~< i ~ n, and count the population of each variety at
each generation. Call N~ the population of variety V/at
time (generation) t. If the fraction of each generation
that dies without leaving offspring is d~ < I, then the
population of variety V~at time t + 1 is:
N~+t = 2(1 -- d=)rauN~ (1)
where mu is the probability that division is not
accompanied by mutation, and the factor 2 accounts
for the division of each organism into two new
individuals.
The above formula assumes that all the offspring of
generation t belong to the same variety. In fact, the
number of individuals of variety V~at time t + 1 will
differ from the above by two terms: the fraction
leaving the variety and the fraction entering it, in both
cases by mutation. Call m~i the probability per unit
time of a mutation from variety V~to variety Vj. These
probabilities are subject to the normalization
condition:
~, mi~ = 1 for every I ~< i <~n (2)
J

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©1978 IPC Business Press Appl. Math. Modelling, 1978, Vol 2, September 201
A m o d e l o f evolution: M . B u n g e
The average number of defectors from variety V/
between times t and t + 1 will be:
D ~ = 2 ~ m o o -- di)N~
A n d the new arrivals to variety V~ in the same time
interval number:
A ~ = 2 ~ mj,(l -- d~)Ni
for they will be drawn from all varieties but Vi. The
factor 2 in D ~ and A ~ shows that we are making the
additional hypothesis that mutants come in pairs--i.e.
if an organism is a m u t a n t so is its sister.
Collecting equations (1), (3) and (4) we obtain the
net average population of variety V~ at time t + 1:
.N[~+I = 2mi,~l -- di)N[ - D i + A i
= 2 [ m i , l - di) - ~ ~ j { 1 - di)]N[
+ 2 ~ ,nj,{1 - - d j ) N l
Because of the normalization condition (2) of the
mutation probabilities, the total probability of a
mutation from V~is:
n~j = ~ rno -- ma = 1 - mu
j~i j 1110, 0111, 1011, 1101, and 1111. The general formula
Introducing this value into equation (5) we are finally
left with:
N~+ ~ = 2(2mu - 1)(1 -- d,)N~
+ 2 ~ mj,O -- d~)Ni
This system of n linear difference equations
summarizes our model of evolution. The solution to
these equations can be obtained as follows. Set:
a =-- i1%tl -- II2(2m. - 1)(1 - d3~oll
b = ltbi~ll = 112mall -- d~)(1 -- 6~j)lt N, =
where f o = 1 if i = j and vanishes otherwise. The
system, (7), can now be condensed into the matrix
difference equation:
N l ÷ 1 = (a + br)N~
where T designates the transposition operation.
Iterating equation (9) we find the general equation:
Nt+k = (a + br)kNt where ke N
The solution to this equation is:
Nt = (a + br)tNo
where No is the column matrix of the initial
populations of the n varieties being considered.
Suppose that all the entries of the evolution matrix
a + b "r are positive and that initially a single variety is
present. Then all the other varieties can emerge
202 Appl. Math. Modelling, 1978, Vol 2, September
simultaneously at the next generation and the
mutation-and-selection process proceeds relentlessly.
Every one of the n varieties prospers once it has
(3)
emerged, and the prosperity of each is conducive to
that of all. (The only qualitative change in this process
can be induced by a sudden environmental change
causing some death rates to jump to unity. ) But, in
(4) general, the process will not be 'explosive':while some
entries of the evolution matrix may be positive, others
may be zero or negative. An overall negative matrix
corresponds of course to extinction.
As long as the death rates di and the mutation
probabilities mu are not specified, the model is a
phenomenological skeleton.
Underlying mechanism
Let us now specify the architecture of our evolving
organisms as well as their mutability (mu) and viability
(d~). We assume that the n varieties of our
biopopulation differ by the composition or
(5) arrangement of a fixed number of modules or basic
units that can be repeated. For the sake of simplicity
we suppose that there are just two kinds of module, to
be called 0 and 1. Thus, if each entity is composed of
four such modules arranged differently, we obtain a
total of n = 24 = 16 varieties, namely 0000, 1000,
(6) 0100, 0010, 0001, 1100, 0110, 0011, 1010, 0101, 1001,
for a variety whose members are composed of m
modules is abc.., m, where each letter is a binary digit.
So much for architecture.
Every change in at least one of the m binary digits
occurring in the formula for a variety is a mutation.
(7) Thus a mutant will differ from its parent organism
either by the total number of O's (or l's) or by the
order of O's and l's. Thus in the above example the
transitions 0000 ~ 1000, 0100 ~ 0110, and 0111 ~ 1111
are mutations and, at the same time, changes in
/v; variety. Of course not all mutations are equally
probable. We shall determine certain relations among
mutation probabilities with the help of the following
considerations on viability.
N~
Considering the environment, we lump all the
- (8) environmental factors into a single, real valued
function e with values in the [ - 1, 1] interval. And we
assume that the probability that an organism
N~ belonging to variety V~survives up to and including
division is symmetrical about the origin. More
precisely, we assume that the death rates di in equation
(1), far from being constant, depend upon the
environmental variable e in the following way:
(9)
~ri(l -- e 2) for -- 1 ~< e ~ 1
1 -- di = ~1 otherwise (12)
(10) where 0 ~< vi ~ 1 for each variety V/.
Since vi is characteristic of each variety and is
environment-independent, we call it the "adaptedness"
(11) of organisms of the variety I'j. (Notice the difference
between this concept and that of fertility. Adaptedness
is a necessary condition for fertility, so the latter is an
indicator of the former.)
The factor (1 - e 2) represents the-selection effected
by the environment. A substantial increase or decrease
in e will have a considerable impact on the fraction
 A model of evolution: M. Bunge
1 -- di of a given population that survives until the A first consequence of the above assumptions is:
next division. In particular, if a catastrophe occurs
(e = + 1), no survivors are left, i.e. di = 1 for all i. On /7/21 ~--- 11131 17/42 ~ / H 4 3
(22)
the other hand if the environmental conditions are /7/14 = 2 m 1 2 m 2 4 m41----- !7114 = 27122171142
optimal (e = 0), then the fraction v~ of individuals of
variety V~survive long enough to divide. The variable e Finally we apply the normalization condition,
is. in sum, the causal or nonstochastic component of equation (6) which, together with the previous
evolution. equalities, yield the mutability matrix in terms of nh 1,
We now introduce an assumption that will allow us m22 and m:
at least to rank the viability values. Our hypothesis is Ilmull =
that the more varied the composition of an organism
the better its chances of survival. Thus in the case
m = 2, where we have only the varieties 00, 01, 10, and "~,'. . . . -,.J -Im..
,,,.-
_-3 ......-., -t
.,3 .,~.- _3,,,~-~_
.....- ~' "~'t4,..... ~- ~ -,,0 !(23)
......
11, we set:

B
~1 -- m2z -- nO m mz2 141 -- mzz -- nO
vl=v,=v f o r 0 0 a n d 11 (13) (1 - nqt)[I --mzz--ml, 1 --m1~ ! --nqt
II 3--mzz - m 3-mzz--m 33-- mzz-- m m t t
Va = v3 = cry, with e > 1, for 01 and 10
We now proceed to trace the evolution of our four-
Finally we relate viability to mutability, namely: we
variety biopopulation. To compute their numbers at
postulate that a mutation is the more probable the less
successive generations we must k n o w their initial
viable its product is. More precisely, we assume:
numbers. Suppose a single .variety is initially present,
m U >1 mik if and only if vj <~ r'k (14) say 1/i, or 00. Call N~ = N its initial population. Then
by equation (7), one generation later, the average
We can now build the matrix of the mutation numbers are:
probabilities in the simple case where rn = 2, whence
n = 4. In this case the viabilities are those indicated in N~ = 2 ( 2 m x x - 1)(1 -- d , ) N
equation (13). Hence the chances of reproducing are:
N2= 2(1--mtl) (1--dl)N
1 - d I = l -- d . = v(1 -- e 2) 3 - - m 2 2 - - #z
(15)
1 - d 2 = 1 - d 3 = :t.t~l -- e 2) (24)
= N,
The hypothesis, equation (14)~ that most mutations are
deleterious, together with the assumption, equation (13), N~ = (l - "22 - m)N~
about the adaptedness values, entails:
If ml 1 ~< ½ or d, = I, the first variety becomes
m41 = m14 ?na2 = m23 = m (16) extinct forthwith. A n d if ml, = 1 while d t < I, Vt
(where this m is not to be confused with the number of prospers without evolving. The necessary and sufficient
kinds of modules per organism, which we have taken condition for the evolutionary process to occur is then:
to be 2), and:
½<m11<1 and 0~<dI<l (25)
m21 = film12 real = fllmaa
(17) If this condition obtains then the lower bound of the
m24 .= f12m42 m 3 4 ----- f 1 2 m 4 3 , population of varieties 1/2 and V3 at first generation is:
with ill, fiE > 1. We-simplify by setting fll = f12 = fl :](1 -- roll)(1 -- da)N
and so are left with:
Since this number equals at least 2, we infer that the
m2l = tirol2 m31 = flmxa
(18) lower b o u n d of the stability of the parent variety is
m24 = tim42 m34 = flm43 given by:
mll I> 3/(1 - dx)N (26)
Simplifying assumptions a n d final e q u a t i o n s Because the upper bound of m , , is I, we infer that for
Since permutations of modules are less radical than a change of variety to occur it is necessary that:
changes in composition, we can make the further
N > 3/1 -- dt (27)
simplifying assumptions that:
For a low death rate of the parent variety, as few as 4
m12 = m13 m24 = m34 (19)
members of it could give rise to the differentiation
Similarly, we posit that the variety O1 is just as stable process.
as 10, and that' O0 and l 1 are equally stable: The above conditions concern only varieties V2 and
V3. For variety V4 to emerge a more severe condition
m22 = ma3 nh, = m44 (20)
must be met, as we see from the last of equations (24),
Our final assumption is that the mutations between the namely, equation (25) plus (m22 + m < I) or,
extreme varieties O0 and 11 proceed via the equivalently, equation (25) plus (m2 x + m24 > 0).
intermediate species O1 and lO, i.e.: Finally we write the general evolutionary equations
(7) for an arbitrary t value. Using the mutation matrix,
II114 = IH1211124 + FH13tH34.
(21) equation (23), we obtain the following System of
m 4 1 = D142Yl121 "~- !Y14317131 first-order linear difference equations:
I~li~IOTHEEK MATHEMATISCH tiff.N~F, UM
L . AMSTiRDAM
Appl. Math. Modelling. 1 9 7 8 . V o l 2. S e p t e m b e r 203
A model of evolution: M. Bungo
N:+a = 2(2mal -- 1)(1 -- dl)N: evolution, equations (7), could be turned, from a mere
descriptive black box, into a mechanismic model
+ (I -- rn22 -- m)(1 -- d2)N2
representing the mutation-selection process.
+ (I -- m22 -- m)(l -- d3)N~
+ 2(1 - m~)(1 - m ~e - m)(1 _ d,)N,"
Discussion
3 -- m22 -- m
O u r model of evolutionary processes is perhaps the
N ? + x = 2(2m22 - 1)(1 - d2)N~ simplest that one can imagine, as it deals with
2(1 -- m11) idealized bacteria assumed to be composed of two
+ (1 - a~)N: units of the same or different kinds. It is a linear
3 -- m22 ~ m
model. Furthermore, it includes a n u m b e r of
simplifying assumptions. A n d yet the final system of
+ 2m(l -- ds)N~ + 2 3 (1 - m11) (1 - d4)N~
m22 m m equations is sufficiently general to invite the use of a
digital computer. This suggests that a more realistic
N~. 1 = 2(2m33 -- 1)(1 - d3)N~ model, say of a plant or animal lineage, must be far
more complicated. However, this should be taken as a
2(1 - roll ) challenge rather than a pretext for refraining from
-(1- d~)~
+ 3 -- m22 -- m modelling evolutionary processes.
The use of our model is presumably methodological
2(1 -- rn 11) rather than substantive. T h a t is, it shows the kind of
+ 2m(1 -- d2)N~2 + .(1 -- d,)N~
~ m22 -- m premises (ddta and specific hypotheses) that must be
added to the extremely general theory of evolution in
N:+z = 2(2m44 -- 1)(1 -- d.,)N:
order to build a precise, albeit highly idealized, model
+ 2(1 -- mix)(1 -- m22 -- m)(l of an evolutionary process. O u r model also shows that
d l)Ntl it is not biospecies but biopopulations (or even entire
3 - m22 - m
communities) that do the evolving. (Species are sets,
+ fl - m 22 - m)(1 - d2)N~ and sets are concepts, not objects that are capable of
changing.) Finally, our model shows that the theory of
+ (1 - - m2z -- m)(1 -- d3)N] evolution, far from-being nonscientific, is a rich matrix
W e shall not a t t e m p t to solve this system. O u r aim for the gestation of any n u m b e r of specific, hence
was only to show how the general equations of precisely testable, scientific theories.

204 Appl. Math. Modelling, 1978, Vol 2, September