10/28/2015

Fundamental Genetics
Lecture 8

Linkage and
Chromosome Mapping
in Eukaryotes

Linked Genes
Genes located in the same
chromosomes
Initiated by Thomas Morgan and
Alfred Sturtevant
Will not segregate independently
Affected by crossing over

John Donnie A. Ramos, Ph.D.

Dept. of Biological Sciences

College of Science
University of Santo Tomas

Linkage vs Independent Assortment

Linked Genes in Drosophila

Red eyes (bw+) dominant to
mutant brown eyes (bw)
Thin wing veins (hv+) dominant to
mutant heavy wing veins (hv)

X-Linked Genes in Drosophila

Cross A
Gray body (y+) dominant to
mutant yellow body (y)
Red eyes (w+) dominant to
mutant white eyes (w)
Cross B
Red eyes (w+) dominant to
mutant white eyes (w)
Normal wings (m+)
dominant to mutant
miniature wings (m)
Due to linkage and crossing
over that occurred during
meiosis

Chromosome Mapping
Determining the distances between genes and the order
of sequence in a chromosome
Uses the frequency of crossing
The shorter the distance between linked genes, the lower the
frequency of crossing-over.
The longer the distance, the higher the frequency of crossing over
to occur.

Frequencies of crossing over:

1. Yellow, white
2. White, Miniature
3. Yellow, miniature

00.5 %
34.0 %
35.4 %

1% of crossing over = 1 map unit (centimorgan, cM)

Distance between linked

genes is related degree of
crossing over

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Distance Affects Crossover

Single Crossover

50% are recombinant gametes

50 % non-crossover gametes
If distance between genes is more than 50 map units,
~100 % crossing over will occur.

Multiple Crossover

Three-Point Mapping
The percentage of crossing over could be used
to map genes in a chromosome
Three criteria needed for successful mapping:
Genotypes of organisms producing the
crossover gametes must be heterozygous
for all gene loci
Cross must be constructed so that the
genotypes of gametes could be determined
based on the phenotypes of the offspring.
Large number of offspring must be
produced

Occurrence of more than one crossovers between non-sister

chromatids.
Produces double crossover (DCO) gametes
If the probability of crossover between A and B is 20% (0.20)
and the probility of crossover between B and C is 30% (0.30),
the frequency of DCO is 6 % (0.06)
Product Law: (0.2)(0.3)=0.06

Determining Gene Sequence

Traits considered:
1. Body color
Gray(y+)
dominant to
yellow (y)

Steps:
Determine 3 possible orders
w-y-ec (y at the middle)
y-ec- w (ec at the middle)
y-w-ec (w at the middle)

2. Eye color
Red eyes (w+)
dominant to
white (w)

Perform a theoretical double cross over

Compare the theoretical DCO with actual DCO (least no.)
Perform theoretical NCOI and NCOII and compare with data

3. Eye shape
Normal (ec+)
dominant to
echinus (ec)

White echinus eyes, gray body

Red normal eyes, yellow body
Yellow body, normal white eyes
Gray body, echinus red eyes
Yellow body, echinus red eyes
10,000

Gray body, white normal eyes

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Unknown Gene Sequence

Unknown Gene Sequence

Total=1109

Observed vs Expected DCO

Not all crossovers can be detected

Observed DCO = double cross-over that actually

occurred
Example: (44 + 42)/1109 = 0.078

Expected DCO = theoretical double crossovers

Product of all the SCOI and SCOII
Example: (82+79+44+42) / 1109 = 0.223
(200+195+44+42) / 1109 = 0.434
DCO exp= (0.223)(0.434) = 0.097

Degree of inaccuracy increases with

increasing distance between linked genes

Coefficient of Coincidence
and Interference
Coefficient of Coincidence (C)
The measure of actual DCOs that occurred
C = Observed DCO / Expected DCO
= 0.078/0.097
= 0.804 or 80.4%

Interference (I)
phenomenon when a crossover event in one region of a
chromosome inhibits a second event to occur in a nearby
region)
I = 1-C
= 1-0.804 = 0.196 or 19.6%
Interpretation: 19.6% of expected DCO did not occur or only
80.4% of expected DCO was observed

Problem 1
A stock of corn homozygous for the recessive linked genes colorless (c), shrunken
(sh), and waxy (wx) was crossed to a stock of homozygous for the dominant wild
type alleles of the genes (+ + +). The F1 plants were then backcrossed to the
homozygous recessive stock. The F2 results were as follows:

Phenotype

Number

Phenotype

Number

+++

17,959

+ + wx

4,455

c sh wx

17,699

c sh +

4,654

+ sh wx

509

+ sh +

20

c++

524

c + wx

12

a.

Determine the distance between the c and sh

b.

Determine the distance between the sh and wx

c.

Determine the distance between c and wx

d.

Give the coefficient of coincidence

e.

Compute for the interference

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Problem 1: Solution
c sh wx

c sh wx

+ + +
+ + +

c sh wx

SCOI
SCOII
DCO

+ + +
c sh wx
+ sh wx
c + +
+ + wx
c sh +
+ sh +
c + wx
Total

=
=
=
=
=
=
=
=
=

c sh wx

+ + +

NCO

Problem 1: Solution

17,959
17,699
509
524
4,455
4,654
20
12
45,832

c sh wx

35,658 = 77.80 %

= 0.0232 or 2.32 %
1,033 = 02.25 %

32

= 00.07 %

= 0.1994 or 19.94 %
Distance between c and wx = 2.32 + 19.94
= 22.26

sh

Problem 2
wx

2.32 mu

Distance between sh and wx = (4466 + 4654 + 20 + 12) / 45832

9,109 = 19.87 %

Problem 1: Solution
c

Distance between c and sh = (509 + 524 + 20 +12) / 45,832

19.94 mu
22.26 mu

In a variety of tomato plant, the mutant genes o (oblate fruit), h (hairy fruit), and c
(compound inflorescence) are all located in chromosome 2. These genes are recessive
to their wild type alleles round fruit, hairless and single inflorescence, respectively.
A
testcross mating of an F1 heterozygote for all three genes resulted in the following
phenotypes:

Phenotypes

Number

Phenotypes

Number

+++

73

++c

348

C = (0.0007) / (0.0232)(0.1984)
= 0.1521 or 15.21%

+h+

+hc

96

o++

110

o+c

oh+

306

ohc

63

I = 1-C
= 1-0.1521
= 0.8479 or 84.79 %

a. Determine the sequence of the 3 genes in chromosome 2

b. Give genotypes of the homozygous parents (P1) used in making the F1 heterozygote.
c. Compute for the map distances between the genes
d. Give the coefficient of coincidence and interference

Problem 2: Solution
+++

ohc

NCO:

ohc
ohc

o h + = 306
+ + c = 348

DCO:

Problem 2: Solution

+h+ = 2

Inference from given data:

+oh

Sequence of genes is not correct

c++

One chromosome contains 2 wild

type alleles while the homolog
contains the 3rd wild type allele

NCO:
DCO:

o+c = 2
SCO I:

Three possible orders of the genes involved:

o h c
o c h
h o c

Find a sequence
that will satisfy
both NCO and
DCO

oh+
++c
+ch
o++
+oh
c++

Satisfies NCO but not DCO

Satisfies DCO but not NCO

SCOII:

oh+
++c
+h+
o+c

ohc

Try if the sequence can

satisfy the SCOs

ohc

(same as + o h)
(same as c + +)
(same as + + h)
(same as c o +)

+++
o h c (same as c o h)
o + + (same as + o +)
+ h c (same as c + h)

= 306
= 348
= 2
= 2
=
=
=
=

73
63
110
96

654 = 0.654 or 65.4%

4 = 0.004 or 0.4%
136 = 0.136 or 13.6%
206 = 0.206 or 20.6%

Total = 1,000

Satisfies both NCO and DCO

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Problem 2: Solution

Problem 2: Solution
c

14 cM

21.0 cM
35 cM

Distance between c and o = (73 + 63 + 2 + 2) / 1,000

= 0.140 or 14 % / cM

C = (0.004) / (0.14)(0.21)
= 0.1361 or 13.61%

Distance between o and h = (110 + 96 + 20 + 12) / 1000

= 0.210 or 21 % / cM
Distance between c and wx = 14 + 21

I = 1-C
= 1-0.1361
= 0.8639 or 86.39 %

= 35 cM