3.

G ENE FL OW

Many models of gene flow. I will cover one model (one-way migration) in lecture. The island
model is covered very well in your text book (p. 48-51). Please read this section carefully.

3.9.a

One - way mi grati on: M igration from l arge l a nd m ass ( C onti ne nt) t o smal l
ne arby i sl and.

Say p and q are the allele frequencies on the island. p* and q* are the frequencies on the
mainland. Every generation a proportion m of the island population originates on the mainland.
After one generation of migration,
p1 = (1-m)p + mp*
p1-p* = (1-m)p + (m-1)p* = (1-m)(p-p*)
p2-p* = (1-m)(p1-p*) = (1-m)(1-m)(p-p*)
pt- p* = (1-m)(pt-1-p*) = (1-m)t(p-p*)
pt = p* + (1-m)t(p-p*)

Q: W hat hap pe ns as t beco m es la r ge i f t he re is no c ount erv aili ng p roc e ss ?

A population of song sparrows on an island is originally monomorphic for a dominant allele (AA).
The island population then begins to receive migrants from the mainland, where sparrows are
monomorphic for a recessive allele a. Every generation, the island population is composed of 5%
migrants. After one generation of migration, what is the frequency of a on the island? After10
gnerations? If the a allele is unopposed by selection on the island, what is the eventual fate of
the a allele?
Q: W hat if ge ne fl o w i s o pp o se d by natur al s elect io n? Co ul d t hi s m ai ntai n
vari ati o n?
A: If the migration rate is low, relative to the strength of selection, migration-selection balance
can maintain genetic variation. Let the relative fitnesses of the genotypes at a diallelic locus be
AA: 1; Aa: 1; and aa: 0.8 on an island population. If we express the fitness of the aa genotype as
0.8=1-s, then the selection coefficient against the recessive homozyote is s . Imagine that the
mainland population is fixed for the a allele and a proportion m of the island population is
comprised of migrants each generation. Thus, migration provides a constant input of a deleterious
alleles into the island population. What is the expected genetic constitution of the island
population with respect to this locus? The eq ui li bri um fre q ue ncy of a rece ssive a lle le
that i s i ntrod uce d i nt o a pop ul ation by mi grati on a nd is op pose d by sel ecti on i s
ap proxim ate ly m p*/s
If 5% of the island population is made up of immigrants, what is the equilibrium frequency of the a
allele on the island?

3.10 R ECO MBINATION

3.10.a

Rec om bi nat ion ge ne rate s va riat ion

Mutation is ultimate source of all variation, but in the short term, a large amount of variation
within population is due to recombination among pre-existing alleles.
The number of possible combinations of genes brought about by recombination is very large.
If an organism has one normal and one mutant gene for each of its 10,000 genes (a reasonable
number; humans may have 30,000 to 40,000 genes), there would be:
210,000
different permutations and combinations of these genes possible in each egg or sperm. This is an
impossibly large number, which essentially means that the variety of gene combinations that can
be created during sexual recombination is infinite!
Although mutation is the only way of producing new variants of single genes, recombination,
brought about through sexual reproduction, produces more new types of individuals much faster
than mutation. In eukaryotic organisms, therefore, recombination is the greatest source of
variation.
Q: If D rosophila m el a nogast er ha s ~ 18,00 0 ge ne s, a nd P = 0.2 ( its act uall y
subst ant ial ly hi ghe r), ho w m any di ffer e nt kinds o f ga met es c ould a n i ndivi d ual
pr od uc e? As s um e t he re a re a m a xim um o f two all ele s at any l oc us.
A:

However, not all gamete types are equally likely, because some of these recombination events
(those between tightly-linked loci) would be rare.
Another example:
Q: You a re i nve sti gat ing a z o ne o f hy bri diz atio n bet we e n t wo po p ulat io ns o f
Lobeli a c a rdi nali s ( ca rd inal fl o wer). T he p o pulati o ns d iffer i n t he av er a ge hei ght of
indivi dual pl a nts ( Po p1= 60c m; Pop 2=40 cm ). F 1 hy brid s a re ex actl y i nt er me di ate in
he ight (50 cm) a nd ar e het er oz y go us at t hree i nd ep e nd e ntly- se gre gati ng l oc i
co ntr oll i ng hei ght ( Ge noty pe r ep re se nt ed a s: + /- ; + /- ; +/- ). Po p1 indivi d ua ls a re
hom oz y go us for the + a lle le at eac h l oc us and Pop 2 i ndivi dual s ar e hom o zy gous for
the a lle le at ea ch l oc us) . Wit h re s pect t o thes e ge nes, ho w m a ny di ffe re nt ki nd s
of ga met e s c an a n F1 i ndiv id ual pr o duce?
Fo r a c hall enge: E ac h + o r al lel e eit he r a dd s or subt ract s 2 c m o f he i ght. Yo u
mat e t wo F1 hy bri ds to p r od uc e t he F2 ge ne rat io n. H o w ma ny d iffer ent ge notyp e s
ar e p o s si ble i n t he F 2 ge ner ati on?

3.10. b

Rec om bi nat ion ca n bre a k up rare gene c om bi nat ions

Recombination breaks up rare gene combinations. If + and alleles affect size at three
independent loci, and the frequency of the + and - alleles at each locus is p=0.9 and q=0.1
respectively, a triply homozygous individual -/-; -/-; -/- will arise occasionally, but will almost
certainly mate with a more common genotype (say +/+; +/+; +/+). So offspring will be
heterozygous at all loci and will be unlikely to themselves have progeny of the -/-; -/-; -/genotype.
Say the triply heterozygous genotype -/+; -/+; -/+ has the highest fitness. Even if two highfitness individuals mate with each other, a large proportion of their offspring will have lower
fitness, recombinant genotypes:
Q: W hat p ro p orti o n o f t he ir o ffs p ri ng wil l hav e t he hi ghe st- fit ne ss ge notyp e a nd
what p ro p ort io n will have l o we r- fit ne s s r ec om bina nt ge noty pe s?
A: