Linkage, Crossing Over and

Chromosome Mapping In
Eukaryotes

Exceptions to Mendelian Principle of

Independent Assortment
W. Bateson, R.C. Punnett sweet pea,
flower color and pollen length
P: red, long x white, short
F1: red, long
F2: 24:1:1:7 ????
Red, long - 583; white, short 170 ; parental
Red, short 26; white, long 24; nonparental

Location of Mendels Genes

Trait

Phenotype

Alleles

Chromosome

Seed form

round-wrinkled

R-r

Seed color

yellow-green

I-i

Pod color

green-yellow

Gp-gp

Pod texture

smooth-wrinkled

V-v

Flower color

purple-white

A-a

Flower location

axial-terminal

Fa-fa

Plant height

tall-dwarf

Le-le

Linked Genes

Thomas Morgan, Alfred Sturtevant, 1911

Cross: two genes located on X-chromosome
y yellow body; y+ - wild type, gray body
w white eyes; w+ - wild type, red eyes
P: yyww x
y+w+/Y
F1 all females wild type, all males mutant
phenotype
F2 98.7% parental phenotypes, 1.3% -yellow
body, red eyes or gray body, white eyes

Chromosome Mapping
The frequency of exchange could be taken as an estimate of
the relative distance between two genes along the
chromosome
Exchange occur due to recombination between loci during
meiosis
Non-parental gametes recombinant gametes
Recombination frequency
RF = # recombinants / #total
Distance between two loci
R = (# recombinants / #total ) x 100 = distance in
cM (now mu map units)

Chromosome Mapping
1% recombination between two genes equals one
map unit centiMorgan, cM
Sturtevant, Bridges, 1923 recombination is not
restricted to X-linked genes
In Drosophila, recombination occur only in
females, not in males
Confirmed the chromosomal theory of inheritance

Single Crossovers
The closer are genes on the chromosome, the less
likely it is that any single crossover event will
occur between them
The farther apart two linked genes are, the more
likely that a random crossover event will occur
At the distance > 50%, linkage cannot be
determined genes behave as unlinked

Crossing-over (CO)
Breakage and rejoining of homologous DNA
double helices
Occurs only between nonsister chromatids at the
same precise place not really
Visible in diplotene as chiasmata
Occurs between linked loci on same chromosome
Cis (coupling): recessive alleles on same homolog
(AB/ab)
Trans (repulsion): recessive alleles on different
homologs (Ab/aB)

Distance Between Two Linked

Genes
Grey and yellow body y+ and y
Wild type and white color of the eyes w +
and w
Cross yy ww x y+w+ Y;
then yy+ ww+ x yw Y
Results: yellow, white 490; wild type 497;
yellow 8; white - 5

Distance between three linked genes

Grey and yellow body y+ and y
Wild type and white color of the eyes w +
and w
Miniature - short wings vs wild type

Genetic Map

Figure 5-4

Copyright 2006 Pearson Prentice Hall, Inc.

Independent assortment
Consequence of independent alignment of
chromosomes in meiotic bivalents
A

A/A ; B/B a/a ; b/b

A/a ; B/b

OR

A;B

A;b

Alternate bivalants

a;B

Alternate bivalents

a;b

Recombination
Creating new combinations of two or more pairs of
alleles
A/A
B/B

meiosis

meiosis

A
B

parental genotypes

a
b

a/a
b/b

A/a
B/b
meiosis

A
B

a
b

parental (P) genotypes

A
b

a
B

recombinant (R) genotypes

Independent assortment
For genes on different (nonhomologous) pairs of
chromosomes, recombinant frequency is always 50%
A/A ; B/B a/a ; b/b

A/A ; b/b a/a ; B/B

A/a ; B/b

A/a ; B/b

A;B

A;b

a;B

a;b

A;B

50%

A;b

recombinants

a;B

a;b

Figure 5-1a

Copyright 2006 Pearson Prentice Hall, Inc.

Figure 5-1b

Copyright 2006 Pearson Prentice Hall, Inc.

Figure 5-1c

Copyright 2006 Pearson Prentice Hall, Inc.

Crossing Over Is a Physical

Exchange Between Chromatids
That crossing over was the result of an
exchange of genetic information between
homologous chromosomes was definitively
shown by the experiments initially
performed by Harriet Creighton and Barbara
McClintock (1931).
They used a combination of genetic and
cytological evidence in their proof.

Crossing Over Is a Physical

Exchange Between Chromatids
Used corn chromosome 9 markers:
c = colorless seed
wx = waxy endosperm

Created a heterozygote with the following

characteristics:
Repulsion configuration of genetic markers
Cytological landmarks on both ends of one
chromosome

Creighton and McClintock

Experiment
Chromosome 9
Large chromosome with 1:2 ratio in arms
Short arm possesses a large knob vs small and
no knob
Long arm carries translocation of chr 8 that
broke the arm near centromere
knob Yg C - Sh Wx translocation parent
Crossed to a plant knobless yg-c-sh-wx-normal

Linkage Problem
In corn, gene C colored seed, c colorless
Gene S full endosperm, s shrunken
A true-breeding colored, full-seeded plant is
crossed with a colorless, shrunken-seeded
F1 - ?
F1 plants test-crossed

Linkage Problem
Colored, full 4,032
colored, shrunken 149
colored, full 152
colorless, shrunken 4, 035
Total
8,368

Linkage Problem
Parental: colored, full 4,032
colored, shrunken
4, 035
Total 8,067
Nonparental: colored, shrunken
149
colorless, full 152
Total 301
Rf: ____________________________________________
If genes in repulsion?