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Author(s): P. Holgate

Source: Journal of Applied Probability, Vol. 3, No. 1, (Jun., 1966), pp. 115-128

Published by: Applied Probability Trust

Stable URL: http://www.jstor.org/stable/3212041

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J. Appl.Prob. 3, 115-128(1966)

Printedin Israel

OF EVOLUTIONARY GENETICS

Smmary

Some comparisons are made between various characteristicsof the genetic

structures of populations of the same size and age, which have (i) evolved

from a small founder population, and (ii) evolved from a population which

has been of constant size throughout the period considered.

1. Introduction

Consider a gene having two alleles A and a. In the absence of selection, and if

there is no mutation, it is possible that one of these will become fixed in a popula-

tion as a result of the random fluctuations in its frequency from generation to

generation. This is generally known as the Sewall Wright effect, and its possible

contribution to evolution has been the subject of considerable discussion. A sum-

mary of Wright's own views and a survey of his mathematical work on the subject

are given in Wright (1964).

Mathematical work on the tendency to homozygosity of a finite population

has been surveyed by Moran (1962), and Kimura (1964). Roughly speaking, for

given initial gene frequencies, the probability that the populations will become

homozygous within a given number of generations may be appreciable for small

populations, but becomes negligible as the population size increases.

Suppose however that a small founder population colonises an area, and in

the course of several generations multiplies to a much greater size. Fixation may

occur at a given locus when the population is small, and the effect of population

growth will then lead to a homozygous population of such a size that fixation

as a result of random fluctuation would be inconceivable in a population which

had always had that same, large size. This phenomenon is known as the founder

principle, associated with Mayr (1942, 1963). Experimental work in relation to it

is reported by Dobzhansky and Pavlovsky (1957), and it has recently been invoked

in the controversy about the origins of geographical variations between snail

colonies in Southern England, (Goodhart 1962, 1963; Cain and Currey 1963a, b).

Despite its interest, little mathematical work has appeared on genetic fluctuations

115

116 P. HOLGATE

selfing populations,in particularthose where the number of offspringof each

individualwas (a) a constant,and (b) a Poissonvariable, and derivedrecurrence

relationsfor the momentgeneratingfunctions,and hencethe meansand variances,

of the numberof heterozygotesin succesivegenerations.He also showed how

the theory could be developedto deal with other systems of inbreeding.Feller

(1951) approachedthe problemfrom the point of view of diffusiontheory and

discussedsome of the difficultiesin formulatingand solving the mathematical

problems.Mode (1964)has consideredanotheraspect.

In the present paper I consider a population which reproducesin discrete,

non-overlappinggenerations.However, instead of keeping the generationsot

constantsize, I have assumedthroughoutmost of the paperthat the population

exactlydoublesin size at each generation,startingwith a singleindividualor pair.

The fact that the overallgrowth is made deterministicin this way bringsabout

two relatedsimplifications.It reducesthe numberof randomlyvariablequantities

and eliminatesthe possibilityof the populationdying out completely.In models

for which this possibilitydoes exist, the interestingresultsare often those con-

ditional on it not occuring,which considerablyincreasesthe mathematicaldif-

ficulty of the problem.In a practicalbiologicalsituation,it would be impossible

for a populationto continuedoublingat each generationindefinitely,but as will

be shown, the effect of the first few generationsof breedingare overwhelming,

and sincein a largepopulationgeneticstructureis not affectedby randomsampling

variancein any case, the resultsobtainedshould have a certainrobustness.

Section 2 is devoted to selfingpopulationsinitiated by a single heterozygote,

and Section 3 to randommating populationsinitiatedby a single pair, each of

whose genes is A or a with probability1/2. From time to time I have compared

characteristics of the structureof the increasingpopulationwiththoseof a station-

ary population on the following basis. Suppose that after t generations,the

growing population has attained a size N. Then for comparisonI consider a

populationwhichhas evolvedthrought generationsof reproduction,maintaining

a constantsize N throughout,havingbeeninitiatedby a highlygeneticallyhetero-

geneous population.Thesecomparisonscan be computednumericallyfor small

values of t, and are given for t = 3, and asymptotic formulae are given for

large t. In Section4, somenumericalresultsaregivenfor thepopulationsresulting

from threegenerationsof breedingwith a fully stochasticmodel.

2. Selfingpopulations

Let us considera populationinitiatedby an Fo consistingof a single hetero-

zygote,Aa.Let each individualof Ft1 contributetwo offspringto Ft, the breeding

systembeing self-fertilisation.Ft will thereforeconsist of 2' individuals.

Oneof the featuresof selfingpopulationsof constantsizeis the rapidelimination

of heterozygotes.(See e.g. Li (1955,p. 103)),and it is thereforeof interestto study

A mathematicalstudy of thefounderprincipleof evolutionarygenetics 117

in Ft then {Zo = 1, Z1, Z2, "*-} is a branching stochastic process of Galton-

Watson type. (For the theory of these processes see Bartlett (1960), Harris (1963)).

The probability generating function of the number of heterozygous offspring

of a heterozygote is

(1) f(s)=

(2+f s).

Since the mean of the distribution given by (1) is 1, the ultimate elimination of

heterozygotes is certain. The p.g.f. of Zt is obtained by functional iteration which

may be carried out explicitly for small t. For instance, the probability that F3

will contain j heterozygotes is the coefficient of sJ in

This distribution is tabulated in the first column of Table I.

The elimination of heterozygotes in a population where each individual con-

tributed a single offspring to the next generation was discussed by Bartlett (1937).

Suppose, for comparison with the above case, that a population had been initiated

by 8 heterozygotes, and had undergone 3 generations of selfing, with each individ-

ual contributing a single offspring to the next generation. The probability that a

given line would remain heterozygous is 1/8, and the p.g.f of the number of

heterozygotes in F3 is therefore

TABLEI

Distribution of number of heterozygotes

after three generations of selfing

Probability

Growing Stationary

Population Population

0 .483459 .343609J

1 .217285 .392696

2 .176514 .196348

3 .077637 .056099

4 .032593 .010018

5 .009277 .001145

6 .002686 .000082

7 .000488 .000003

8 .000061 .000000

118 P. HOLGATE

probabilityof complete eliminationof heterozygotes,and the probabilitythat

theirnumberwill be large,are greaterfor the growingpopulation. The variances

are 1.5 for the growingpopulationand 0.875 for the stationaryone.

The varianceof Z, for generalt may be obtainedby lettingf -2 in Formula(7a)

in Bartlett(1937), or by substitutingfor a2 in Formula(5.3)in Harris(1963,p. 6),

eitherof whichlead to:

by the asymptoticformulaof Kolmogorov(1938)as

2 4-

(5) Pr(Z, =0) 1- tf(1)

I

t

1

In fact, the p.g.f. (1) and result(5) are mentionedby Kolmogorovas an example

at the end of Section4 of his paper,wherethey arisein the solutionof a different

problemin populationgenetics.It may be noted that t has to be quitelargefor (5)

to give good numericalresults. For t = 40 when (5) gives 0.9, the correctvalue

obtainedby iterationis 0.9142.Howeverthe derivationgivenby Harris(1963,p. 20

ff.) suggestst/(4 + t) as a betterapproximation,and for t = 40 it gives0.9091.For

comparisonconsider an Ft which had arisen from an Fo of 2t heterozygotes,

evolving throught t generationsof selfing at constant population size, and let

Zt be the numberof heterozygotesin Ft in this population.(It should be borne

in mind that the sequence{Z'}, althougha stochasticprocessin the mathematical

sense, does not representthe evolution of a definitepopulation.) By reasoning

similarto that given in the case t = 3, the distributionof Zt is binomial with

n = 2t, p = (1/2)t, and hence

Comparisonof the variancesgivenin (4) and (6), and the probabilitiesthat hetero-

zygotes will be absent, given in (5) and (7), togetherwith the numericalresults

of Table I, illustratethe fact that selfingpopulationswhichhave reacheda large

size by repeateddoubling will be more variable in respect to proportions of

heterozygotes,and in particularwill be more likely to contain none, than those

studyof thefounderprincipleof evolutionary

A mathematical genetics 119

of the same size and age that have always been that size. The stochastic process

describing the number of individuals in Ft which are homozygous for a given

allele, say A, is not a branching process, nor even a Markov process. However,

the probability P, that Ft will consist entirely of AA's, has a simple recurrence

relation. The probabilities that the offspring of a heterozygote are (AA,AA),

(AA,Aa) or (Aa,Aa) are 1/16,1/4,1/4 respectively. It is then required that the

offspring should consist entirely of AA's after (t- 1) generations, which leads to

11 1 2

Pt =- + Pt-1 + Pt2-1

the probability that the population ultimately consists entirely of AA's satisfies

11 1 1 2

P=- +-p +P2

16 4 4

of which the relevant root is P = 1 1/2 - V2 = 0.085786. For the comparable

stationary population the chance that all its members are AA is given by

(1/4 + 1/2 * 1/4 + 1/2 * 1/2 * 1/4)8 = 0.001342.

In the doubling population the elimination of heterozygotes occurs with

probability one. When it has occurred the population which will then consist

entirely of AA's and aa's will continue to increase, but with the numbers of these

types, and hence of the two alleles, in a constant proportion. The remainder

of this section is devoted to obtaining some properties of Y,the random proportion

of AA's when the heterozygotes have been eliminated. Clearly, Y can take only

those values whose expression as a binary fraction can be written in terminating

form, and every such value in the interval [0,1] is taken with positive probability.

It is therefore an example of a random variable whose distribution function has

jumps at a set of points everywhere dense (in [0,1]), although this peculiarity is a

mathematical consequence of insisting on the population doubling exactly at each

generation. If Yt is the proportion of the limiting population belonging to lines

which became AA at exactly the tth generation, then Y = t 1 Yt.It is interesting

to note that the stochastic process

2fYt, j=1,2,-.,

t= I

is such that almost every realisation is monotonically increasing, the form of the

'remainder' term in the submartingale decomposition theorem (Loeve, 1963,

p. 389). Some properties of Y may be obtained by enumerating the possible

structures of F1, and equating the unconditional expectations of certain variables,

to the appropriate expectations obtained after conditioning by the outcome of F1.

The possible F1 's are listed in column 2 of Table II, preceded by their probabilities.

120 P. HOLGATE

Column 3 gives the conditionalexpectationof Y, columns 4 and 5 give thecon-

ditional varianceof Y and the probabilitythat Y = 0, wherev and P denotethe

unconditionalvalues of these quantities.Column 6 gives the conditionalexpec-

tation of ez, wherem(z) is the unconditionalvalue.

TABLEII

Probabilitiesof possibleF1's, and values of variouscharacteristicsof population

structureconditional on these, for the growing,selfingpopulation

1

AA,AA 1 0 0 ez

16

- AA,Aa 4 ^ 0 e '2m( z)

1

AA,aa 0 0 ezI

2

Aa,Aa 2 \ (z

AA

aa,aa 0 0 1 1

16

secondmomentabout zero:

414+)14(16+4)

6 +4v+8(1+

-v+4T6H+4

which v = For the

gives of the eliminationof

1/12. probability A,

1 1 1 2

P=- +-P +p-p2

moment generatingfunction written down from the last column simplifiesto

A mathematicalstudy of thefounderprincipleof evolutionarygenetics 121

2z for convenience, +(z) is found to satisfy the functional relation

Since Y is bounded, all its moments exist, and expansion of the m.g.f. in (9)

provides a recurrencerelation for the even order central moments. The first few are

1 11 121 61663

#2 2= , P4 =

720 6 = 36288' P8 79315200'

of which the first value confirms the solution of (8). Since the probabilities of

the end points are known exactly, the moments of the distribution conditional

on fixation not occurring can be calculated from the formula

The coefficient of kurtosis of the conditional distribution is f2 = 2.3071.

3. Random mating

Notation used in this section is not everywherecomparable with that of the last.

Since a random mating population cannot be initiated by a single individual,

in order to make the situations in this section and the last as similar as possible,

I have assumed that the population begins with an F1 consisting of 2 animals,

each of whose 4 genes are A or a with probability 1/2. This means that if the

mathematical definition were extrapolated backwards, independently of its

interpretation, we could imagine an Fo consisting of a single heterozygote. As in

studies of random mating populations of constant size, only the number of genes

in each generation is considered, and they are supposed to be determined inde-

pendently, being A or a with probabilities equal to the proportions in the previous

generation. Moran (1962, pp. 12-20) has discussed the implications for the form

of the offspring distribution of the constant population model and analogous

considerations apply here. Let Zt be the number of A genes in F,. The distribution

of Zt conditional on Z- = z,_ is binomial with n =2'+ p = (1/2)tzti.

The distribution may be computed explicitly for small t, and for t = 3 it is given

in the first column of Table III. For a comparable population of constant size,

suppose that Fo had consisted of 8 heterozygotes, which had then evolved through

3 generations of random mating. The distribution of Z3, the number of A's in

F3 is given by post-multiplying the 1 x 17 row vector with 1 in its ninth column

and zeros elsewhere, three times by the matrix (ail) with elements

122 P. HOLGATE

1

iJ

al= ( 16- ( i

{ HJ) (6) 16)

(Feller, 1951). The resulting distribution is given in the second column of Table III.

In this case the probability of elimination of A can be seen to be greater for the

growing population. The variances of the proportion of A genes are 0.0962 and

0.0440 for the growing and stationary populations respectively.

TABLEIII

Distribution of number of A's after three generations of random mating

(The distribution is symmetric about 8.)

Probability

Growing Stationary

j Population Population

0 .0996 .0068

1 .0284 .0155

2 .0423 .0290

3 .0505 .0457

4 .0562 .0639

5 .0607 .0816

6 .0738 .0964

7 .0655 .1062

8 .0661 .1096

of A genes in successive generations.

It will be shown that Yttends to a limit Y as t -+ oo, and some of the properties

of Y obtained.

A recurrence relation can be derived for the second moments of the Zt about

the origin as follows: let P(z,t) = Pr(Zt = z),

2t"I

He(t) = 2 x2P(x, t)

=O 0

t 2

=x y(2X

= ]

P(y,t - 1) X2y

y=0

2= P(yt - 1) 4(2

2y 1I1)

+ - --

2(H; -)+( 2-^l )82( -)

_

The step from the third to the fourth line follows from the fact that the second

summationis the second moment about zero of a binomial distributionwith

n = 2+, p = y/2t. On writing 4 = u2 + ,u2 a recurrencerelationfor the var-

iancesof Z, is obtained

82(t) = - 1 2(t-1) + 2t-.

(4

tion

(11) v2(t)= v2(t- 1) +

V-4T ~- v2(t 1) )

Since the variancesof all distributionson [0,1] are boundedabove by 1/4, (11

shows that the sequence tends monotonically, as t -> oo to a limit 1/4.

On writing(11) in the form

I I 1

( V2(t)) ()(--2(t-))

an explicit solution can be obtainedfor any t,

More lengthy algebraleads to the following recurrencerelation for the fourth

momentsv4(t)

{/IV 1 I 2t 3 1\t

V4(t) = )

4(t()

d2) +

(,-,){~) )_ -() ) T ).

Withinitial values v2(0) = v4(0)=0, computationusing (11) and (12) leads to the

limits

(13) v2 = 0.105606, V4= 0.020182.

For everyk,

V2k(t) = ( Yt - )

t

-tPr (Yt-1 = q) 2 Yt-=

= V20(t-1),

124 P. HOLGATE

then using the symmetryof Yt-1 about ?. Hencethe sequencev2k(t) is increasing

and it is bounded above by (1/2)2k.Hencethe momentsequencesof the Yttend to

a limiting sequenceand since this must be the moment sequenceof a bounded

random variable,it defines it uniquely, and the convergenceof the sequence

{Yt} follows from the second limit theorem of the probabilitycalculus. (See

e.g. Rao and Kendall (1950)).Alternatively,{&Yt| Yt_-=I Yt-} = Yt-1 and so

{Yt} is a martingale,and the fact that its random variables are bounded is

easily enough seen to ensureconvergenceby the martingaleconvergencetheorem

(Doob, 1953,Chapter7).

In this paragraphand the next I obtain upperand lower boundsfor the proba-

bility of ultimatehomozygosity,showingin particularthat it is less than one. In

the presentproblem, actual computationfor the early generationsfollowed by

numerical extrapolation enables the exact value to be calculatedaccurately,

but the methods of obtainingthe bounds are of interest,and may be of value in

parallel studies, since it is clearly importantto know whetherthe probability

of absorption is bounded away from one. The coefficientof kurtosis of Y is

P2 = 1.809738.Hence using the generalisationof Tchebychef'sinequalityquoted

e.g. by Cram6r(1946,p. 256)

Pr Pry( | - 1.5385997)

= 0.3022.

The probabilitythat a given allele, say A, will be fixed is less than half this, i.e.

0.1511.

Let yt be the probabilitythat the populationbecomeshomozygousat exactly

the tth generationconditionalon it being heterozygousat the (t - 1)th, y, the

probability that it becomes homozygous at or before the tth generation,and

y = lim, . Vtthat it ultimatelybecomes homozygous. The chance that fixation

will occur precisely at time t can be shown to be greatestwhen Ft-_ contains

only one representativeof one of the alleles,and least when they are presentin

equal numbers.Hence

1 )t 2t (1 )t 2t+1

^(~~~~~-a'^ ))"-

A mathematicalstudy of thefounderprincipleof evolutionarygenetics 125

k+

>=Y + -)

[1

+(l-

(1 )k+i

_- 1 )\ (

e-)

(14) ( 1 k+2

Y-k + (Y) [ ) 1 1 -

([2eI)}]

generationhavebeenobtainedup to k = 5, and aregivenin TableIV.

TABLEIV

Probabilitiesof the fixation of A after t generationsof randommating, for the

growingpopulation

k 1 2 3 4 5

1 y > 0.105598,

(Aitken, 1926) a numerical approximationto ly may be obtained from the

terms given in Table IV. Applying the method to I72, i73 and I74 gives

Iy = 0.10669, and applying it to I73, I74 and Iy5 gives Iy = 0.10672, which

is thereforeprobably correct to 5 decimal places. Acceptingthis the moments

of Y conditionalon absorptionnot having taken place may be calculatedfrom

(10) and (13),

t2 = 0.06643, r/4 = 0.00982, P2 = 2.23.

random mating at constant size, the probabilityof fixation is approximately

1-(1-(1/2)t+1)' (Moran, 1962, Chapter 4), and this approaches zero very

quickly as t increases.

126 P. HOLGATE

4. A fullystochasticmodel

A simplefully stochasticmodel is obtainedby lettingthe numberof offspring

of each individual be, not a constant, but a Poisson variable. Bartlett (1937,

Formula(7b))obtainedthe variancesof the numbersof heterozygotesin successive

generations,(without conditioning on the population as a whole remaining

non-extinct).For the case where(i) the Poissonmeanis unityand hencethe mean

populationsize is constant,(ii) the numberof generationssufficientlylargefor the

probabilitythat the heterozygousoffspringof a given individualwill not have

been eliminatedto be given by the asympoticformula derivede.g. by Harris

(1963, p. 18),Formula 9.5), and (iii) the founder population so large that the

chance of completeextinctionis negligible,Bartlettalso showedhow the proba-

bility that the population should consist entirely of homozygotes could be

calculated.

In the remainderof this sectionI presentsome numericalvalues relatingto the

results of three generationsof mating,for a populationinitiatedby a single in-

dividualand havinga mean rate of increaseof 2, and for a populationinitiated

by 8 individuals,and having a mean rate of increaseof unity. If the numberof

offspringof each individualis a Poisson variablewith mean 2, and the offspring

of a heterozygoteare AA, Aa, aa with probabilities1/4, 1/2, 1/4. The trivariate

stochasticprocessgivingthe numbersof the threetypesin successivegenerationsis

a multi-type Galton-Watson process (Harris, 1963, Ch. 2) with generating

functions

f2(sl,s2,s3) = exp2 s + + 3-

multivariatefunctionaliteration.Fromit, the probabilitiesthat variouscombina-

tions of classes will have no representativesmay be computed.The probability

of completeextinctionturnsout to be 0.192975.On multiplyingup the remaining

probabilities,the probabilitythat a population,given that it is still in existence

after three generations,consists entirelyof AA's, or entirely of homozygotes,

may be calculated.The comparablestationarypopulationis obtainedby taking

the mean number of offspringas one instead of two, and startingwith eight

heterozygotes.In this case the chanceof total extinctionin F3 is much smaller,

0.023558.In Table V a numberof probabilitiesrelatingto the F3's of various

models consideredare collected together.

A mathematicalstudyof thefounderprincipleof evolutionarygenetics 127

Acknowledgements

I am gratefulto Mr. J. G. Skellamfor severalhelpfulsuggestionsmade in the

course of this work, and to Mr. K. Lakhaniand Mr. D. Spaldingfor assistance

in computation.

TABLE V

stochastic growth models probabilities are conditionalon complete extinction

not having occurred.

Probabilitythat F3 consistsentirelyof:

(i) AA's (ii) homozygotes

Growingpopulation

Deterministicreproduction 0.0838 0.4835

Stochasticreproduction 0.1580 0.4555

Stationarypopulation

Deterministicreproduction 0.0013 0.3436

Stochasticreproduction 0.0645 0.3222

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(1936b) The causes of area effects. Heredity 18, 466-471.

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