,

.
-
.
.

.
2 ( - - )

.
4 4.

.
4:4 , ( .
) .

4 .
4 .
.

2:2:2:2
.


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 .

6
The
8
four types of asci are produced in equal proportions.
 2:4:2
.
4

- .
.


.
4:4
 Recombination frequencies of pairs of genes
reflect the distances between them along a
chromosome

Think of DNA as long, linear molecule

Genes arranged along length
Crossover can occur anywhere along length
with equal likelihood

The probability of 2 genes crossing over and

undergoing genetic recombination increases
with the distance separating them.

13
 ,.

, , ,
.

( )map units = m.u.

)c.m.( centimorgan
, .
 50 m.u.
( ) 2 .

) - Parental Type (PT

) Recombinant Type (RT .
) Double Crossing Over (DCO
.
 AaBb X aabb

A B a b
P: X
a b a b

A b a B
A B a b
F1: ; ;
; a b a b
a b a b

A B a b
P: X X
a b a b

A B a b A b a B
F1: ; ;
a ; a b a b
b a b
 :
AaBb .aabb
:

54 aaBb
46 - Aabb
458 - aabb
442 AaBb
.
:

P.T. !
:
 :
:

10 .
.1 ?AaBb
:
.1 10 , 10% -
%90 - AaBb, ( P.T.
,)aabb . , 45 %
. AaBb
.2 ?Aabb
:
.2 Aabb 10 % .
, - 5 %
( 5% ) aaBb

(
):
 :

;

:
AaBbCc x aabbcc

. "
( )P.T. (.)D.C.O
:

:
(P.T. ) .AbC, aBc
( D.C.O. ) .abc, ABC

( )m.u. :

X 100
 ( ) m.u. :
X 100

:
,
.

, :
,
,
.

= ( ) .1
 :
III .B, V, G :

:

vBg
VbG
Vbg
vBG
VBg
vbG
VBG
vbg
, :
. :
 :

. :R. T.

( P.T. ) ,
,
( R.T .)Vb
.

. :
 .2 b G- .28 m.u.

.3 V( G- ):

:
)R.T. (VB
)R.T. (BG
D.C.O. 2 !

( : B G- B V-)18 + 28 = 46 :

. :
D.C.O. , ,:
0.28 x 0.18 x 1000 ~ 50 D.C.O.
( )
)22 + 18( 40
, ,40/50 = 0.8 : .1-0.8 = 0.2 :
20 % !
Recombination data for 3 sex-linked genes, echinus, scute, crossveinless
P1: ec sc cv x + + + The
ec sc cv
ec sc cv distance
F1: + + + x ec sc cv between sc
Gametes from F1 F2 F2 Genotypes Observed and cv is
Female Phenotypes Number 20%
Male Female )9.1+10.9(
parental combinations
Conclusion
echinus scute ec sc cv * 934 2108 X 100 = 80.0% The gene ec
ec sc cv ec sc cv ec sc cv
crossveinless 2635 is in the
+ + +
wild type
+ + + + + + *1174 Nonrecombinants middle
ec sc cv
new combinations
between ec and sc
(1st
scute
+ sc + + sc + 140 239 X 100 = 9.1% approxima-
+ sc + sc ec cv 2635 tion)
ec + cv echinus ec + cv ec + cv 99 recombinants ec-sc
crossveinless sc ec cv new
new combinations combinations
between ec and cv between
sc and cv
crossveinless + + cv + + cv 124 288 X 100 = 10.9%
+ + cv sc ec cv
2635
echinus scute ec sc + 164
ec sc + ec sc + sc ec cv recombinants ec-cv

Since the viability of the triple mutant is less than the *

w.t, we use two mutation in cis and the third in trans.
Recombination data for 3 sex-linked genes, echinus, scute, crossveinless
sc + cv The distance
P1: sc cv X + ec +
sc +
+
cv between
F1: + ec + X sc ec cv sc and cv is
Gametes from F1 F2 F2 Genotypes Observed 16.8%
Female Phenotypes Number )6.6+10.2(
Male Female
parental combinations
The values are
scute
sc + cv sc + cv 1828 more reliable
3607 X 100 = 83.2%
sc + cv crossveinless sc ec cv 4338
echinus + ec + 1779 nonrecombinants Why there are
+ ec + + ec + sc ec cv no double
new combinations crossovers?
between sc and ec
scute echinus
sc ec + sc ec + 148 287 X 100 = 6.6% We probably
sc ec + sc ec cv 4338 need
+ + cv crossveinless + + cv + + cv 139 recombinants sc-ec Larger number
sc ec cv
of progeny
new combinations
between ec and cv
new
echinus + ec cv 227 combinations
+ ec cv + ec cv 444 X 100 = 10.2%
crossveinless sc ec cv
4338 between
sc and cv
scute sc + + 217 recombinants ec-cv
sc + + sc + + sc ec cv
 Double Crossovers

A-B-C:
1 2
A B C a b c
x
a b c a b c

ABC
no c.o: Parental
abc 2nd approximation

Abc A B C ABc A B C
c.o. in #1: c.o. in #2:
aBC a b c abC a b c

d.c.o. : AbC A B C
(double) aBc a b c
Table 17-3: Recombination data for 3 sex-linked genes, echinus, scute, crossveinless

F1: sc + cv
+ ec + X sc ec cv The gene in the
F2 Phenotypes F2 Genotypes Observed middle is
Number separating from
Male Female the outer genes as
a result of
echinus + ec + + ec +
sc ec cv 8576 17384 X 100 = 83.64% d.c.o
20785
scute
sc + cv sc + cv
sc ec cv
8808
non-recombinants
And the frequency
crossveinless
of this group is
the lowest
scute echinus sc ec + sc ec + 681 1397
sc ec cv 20785 X 100 = 6.72%
)#1(
crossveinless + + cv + + cv 716 recombinants sc-ec
sc ec cv
only
echinus + ec cv + ec cv 1002 1999
crossveinless sc ec cv 20785 X 100 = 9.62% + ec +
)#2(
scute sc + + 997 recombinants ec-cv
sc + + sc ec cv sc + cv
only
#1 #2
scute echinus sc ec cv sc ec cv 4 5
crossveinless sc ec cv 20785 X 100 = 0.02%
)#1+#2)
wild type + + + 1 double crossovers
+ + +
sc ec cv sc-ec-cv
map distance sc-ec = 6.72 + 0.02 = 6.74
map distance sc-cv = 6.72 + 9.62 + 2x(0.02) = 16.38
map distance ec-cv = 9.62 + 0.02 = 9.64
 Summary of mapping using 3-point linkage analysis
1. Determining the distances between each gene pair.
2. Determining the order of the genes
3. Determining the d.c.o group
4. This group will be counted twice, once with c.c. #1 and once with c.o. #2
5. The distance (ac) = (bc) + (ac) .

BUT, d.c.o. is rare than the expected!!!

If there is no dependence between c.o. #1 and c.o. #2 ,THAN:
f(d.c.o.) = f(c.o. #1) x f(c.o. #2)
In the example above:

c.o. #1 = 0.0672, c.o. #2 = 0.0962 ,

SO: (d.c.o.)exp = 0.0672 x 0.0962 = 0.0065 ,

BUT (d.c.o.)obs = = 0.0002 (0.02%)

Mller: (d.c.o.)obs / (d.c.o.)exp = coincidence coefficient

c.c. = 0.0002/0.0065 = 0.03, Meaning only 3% d.c.o. of the

expected did occur !

Lets define Interference = 1-c.c. ,I = 0.97

Different Interference for different distances.

Map unit: 1 morgan = 1% recombinants

How can there be maps of over 50 morgan units? + ec +
sc + cv
#1 #2

sc
6.7
ec
9.6
cv
8.4
ct
16.0
v
11.1
g
If there is no genetic marker
0.0 6.7 16.3 24.7 40.7 51.8 ...
between the external genes
+ +
sc cv
#1 #2

We will get a lower estimation

of the real distance and
as the distance is longer
the deviation is larger

syntenic genes: genes linked on the same chromosome

(a) double crossovers when gene order is
A-B-C
A B C A B C A B C parental combinations
(noncrossover
chromatids)
a b c double crossovers
A b C classes (AbC,
a b c a b c a B c aBc)

The middle gene is moving

(b) double crossovers when gene order is from cis to trans
B-A-C
B A C B A C B A C
parental
b a c combinations
B a C double crossovers
classes (aBC,
b a c b a c b A c
Abc)
(c) double crossovers when gene order is
A-C-B
A C B A C B A C B
parental
a c b combinations
A c B double crossovers
classes (ABc,
a c b a c b a C b
abC)
 15 20 In this example, what are the results we are
A B C expecting if we count 200 progeny?
(d.c.o.)exp = (0.15)x(0.20)x200 = 6
The distance AC

1. IF I = 0.3 ; C.O.C= 70%

2. (d.c.o.)obs = (d.c.o.)expxC.O.C. = 6x0.7 4

2 AbC
2 aBc
13 Abc
67 ABC A-B: (0.15) 200 4 = 26
Parental: 200 (4+26+36) = 134 13 aBC
67 abc
18 ABc
B-C: (0.20) 200 4 = 36
18 abC