Lecture 9

BIOS 3100 Spring 2025

Chapter 4 – Gene Linkage and Recombination
 Genes on
different
chromosomes
assort
independently
R/r; Y/y x rr; yy

RY Ry rY ry

ry RrYy Rryy rrYy rryy

1 :1 :1 :1

Parental
Recombinants
Bateson and Punnett
Experiment

Expected 9:3:3:1
215 71
71 24
Parental Found
Recombinants MORE Less Less MORE
Linked alleles tend to be inherited together
Linked genes
pr vg / pr vg x pr+ vg+ / pr+ vg+

Gametes

pr vg

pr vg
Meiosis
pr+ vg+
No crossover

pr+ vg+ All parental classes

Crossing over produces new allelic
combinations

F1 F1 gametes

Recombinant Classes
chiasma
Chiasmata are the sites
of crossing over
between homologous
chromatids
 Crossing Over
• Crossing over is the exchange of DNA between non-sister
chromatids to form recombinants
• Occurs during prophase I of meiosis
• Does not occur in the meiocytes of fly males and is less
frequent in human male meiocytes
• The rate of crossing over is proportional to the distance
between genes
• Appearance of non-parental types (recombinants) is
<50%
• Multiple chiasmata can form between chromatids or
with different chromatids
 Linkage Terminology
• Linked or not?
• Linked AB/ab
• Not linked (different chromosomes)  A/a; B/b
• Linkage unknown Aa.Bb

• Allele conformation in dihybrids

• Cis conformation – Dominant or wt genes on same homolog
(AB/ab, ++/ab)
• Trans conformation –Dominant of wt genes opposite
homolog (Ab/aB or +b/a+)
 How do we test for
linkage?
Linked Genes Unlinked Genes

male

Recombinants < 50% of progeny Recombinants = 50% of progeny

Drosophila melanogaster has four
linkage groups
 Distance between two linked
genes is determined by
recombination frequency

•One genetic map unit (m.u.) is the distance between genes that gives 1/100
recombinants. Sturtevant, 1911.
•Recombination frequency (RF) - # of both recombinant classes/total progeny
•Genetic map unit = RF x 100
•Map units are also known as centimorgans (cM)
•Map units are not a physical distance but a hypothetical distance
Map distances are generally additive

What additional experiment do we have to do

to determine which map is correct?
Chapter 4: Question 9.1

Three linked genes (A, B, C) on a chromosome show the

following recombination profiles between them:
Genes A and B: 20% recombinants
Genes A and C: 5% recombinants
Genes B and C: 15% recombinants

From these recombination measurements, we can best conclude:

a. that the gene loci must be positioned on the chromosome as in

the adjacent figure (relative to the centromere and telomeres).

b. that the gene loci must be positioned in the chromosome as in the

adjacent figure (relative to the centromere and telomeres).

c. the relative order of the genes, but not their position in the
chromosome
Often, scientists set up crosses with three mutations
to create linkage maps

vermilion eye color (v) cut wing (ct) crossveinless (cv)

WT eye color (brick red) WT wing

Starting materials: cv and ct double mutant strain
v mutant strain

Experiment and Observation

P: cv and ct mutant v mutant

v+/v+ . cv / cv . ct / ct v/v . cv+ / cv+ . ct+ / ct+

F1: F1 females tester males

v/v+ . cv / cv+ . ct / ct+ v/v . cv / cv . ct / ct

F2 Analysis Observation in F1: all progeny are WT

Interpretation: recessive mutations
The cv and ct mutations are autosomal.
 Not Linked: # of parental = # of recombinant
F2 Analysis
Linked: # of parental > # of recombinant
Describe a linkage relationship of these three genes.
If linked, calculate genetic distances and make a genetic map.
Parental: v+ . cv . ct and v . cv+ . ct+
Magic strip
Magic strip

Magic strip

R R
R R
R R
R R
R R
R R
1448 R=268 R=191 R=93
P = 1180 P = 1257 P = 1355
Conclusion: These three genes are linked.
RF (v-cv) = 268/1448 RF(v-ct) = 191/1448 RF(cv-ct) = 93/1448
 RF (v-cv) = 268/1448 = 0.185  18.5 cM
RF(v-ct) = 191/1448 = 0.132  13.2 cM
RF(cv-ct) = 93/1448 = 0.064  6.4 cM

Construct a genetic map (relative locations of genes on one chromosome).

18.5 cM

v cv

13.2 cM
ct
6.4 cM

13.2 + 6.4 = 19.6

Where does this discrepancy (18.5 vs. 19.6) come from?

 Double recombinants arising from two crossovers

Magic strip
Correct gene order after mapping experiment

Parental: v+ . cv . ct and v . cv+ . ct+ --- Original order

These double recombinants were categorized to the parental classes.

R
R
R
R
R
R

RF(v-cv) = 276/1448 = 0.191 = 19.1 cM

 19.1 cM is not equal to 19.6 cM (13.2 + 6.4).

What is the theoretical double recombination frequency in this case?

Expected: [0.132 (RF v-ct) x 0.064 (RF ct-cv)] = 0.008

0.008 x 1448 (total F2) = 12.23 double recombinants

Observed: 5 + 3 = 8
This is due to interference – a crossover in one region inhibits a crossover
in an adjacent region

19.1 cM

v cv

13.2 cM
ct
6.4 cM
A true breeding fly with two mutations, one for white eyes and the other causing
causing miniature wings, was crossed with a wt fly and the result shown below.
All the F1 progeny were wild type. From these results what can you conclude?
 Learning Outcomes
After studying the material in this lecture, students
should be able to:
• Use recombination frequency to determine if genes
are linked on the same chromosome
• Use recombination frequency in a test cross to
calculate the chromosomal distance between genes
in map units
• Determine the order and distance of genes when
testcrosses are performed with three genes