QUESTIONS AND PROBLEMS 3

1. Define the following terms:

Chromosome-mediated transfer
Genetic distance
Heterokaryon
Hybrid cell
Interference (positive vs negative)
Linkage group
Linkage map
Liked genes
Ordered tetrad
Somatic cell hybridization
Syntenic genes
Unordered tetrad
2. Suppose that all the gene loci in particular organism map into seven separate linkage
groups. How many chromosomes are present in the somatic cells of this species?
3. Criticize the following statements:
(a) Crossing over results in the formation of recombinant type.
(b) Genes that are present on the same chromosome fail to assort independently
(c) A genetic map shows the dimensional relationship of genes on a chromosome
(d) The farther apart two genes are on a chromosome, the greater is their frequency of
recombination.
4. Consider the following linkage data:
Gene loci: a-b
b-c c-d d-e
c-e
Map distance: 8
6
2
4
6
Construct a chromosome map of these loci
5. Albinism in mice can arise as the homozygous expression of one or the other (or both)
of two recessive genes, c and d. Individual must have the dominant alleles of both
genes ( C-D-) to express normal color. Suppose that mating between a dihybrid male
(CcDd) and females of a doubly recessive albino strain (ccdd) produce 148 albino and
52 colored offspring. Are the two loci linked?
6. In tomatoes, the gene for round fruit shape (O) is dominant to its allele for elongate
shape (o), and the gene for smooth fruit skin (P) is dominant to its allele for peach (p).
Two series of crosses involving these genes produced the following results:
Cross 1: P: round, smooth X elongate, peach
F1: all round, smooth
Cross 2: P: round, peach
X elongate, smooth
F1: all round, smooth
When the F1 of Cross 1 were test crossed, they produce offspring in the following
proportions: 46% round, smooth; 4% round, peach; 4% elongate, smooth; 46%
elongate, peach. In contrast, testcrosses involving the F1 of Cross 2 gave the following
result: 4% round, smooth; 46% round, peach; 46% elongate, smooth; and 4% elongate,
peach. Explain these results, designating the genotypes of all individuals concerned.
7. The following data were obtained from a series of two-factor testcrosses. Explain these
results in term of the relative locations of the different genes on linkage map.
Gene loci : a-b b-c a-c b-d a-e d-e a-d
% Recomb: 42 44 50 48 50 50 50

8. In rabbits, genes are two loci, b and c, interact to produce coat color; B-C- individuals
are black, bbC- individuals are brown, and cc (B-cc or bbcc) individuals are albino.
Certain crossing data suggest that these are loci are linked. In one series of
testcrosses, mating between a dihybrid male and female of the doubly recessive
albino strain resulted in 65 black, 34 brown, and 101 albino offspring. (a) Are the genes
linked in the cis or trans arrangement in the dihybrid parent? (b) Calculate the map
distance between these loci. (c) The dihybrid male in this series of testcross was the
product of a mating between members of two homozygous strains. What were the
genotypes of these homozygous strain?
9. Nail-patella syndrome in human is characterized by congenital abnormalities of the
finger nails (and sometime toenails) and of the patellae (kneecaps). The gene for this
disorder is dominant and is located in chromosome 9 about 10 map units away from
the ABO locus. Suppose that a man with nail-patella syndrome and type A blood
marries a normal woman with B blood. The mothers of both the husband and wife are
normal and have blood type O. (a) The husband and wife have two children. One with
type A blood and the other with type B blood. What is the probability that both children
are normal (that is, do not have nail-patella syndrome)? (b) The couple are now
expecting another child. What is the chance that the child will have nail-patella
syndrome and O blood? (c) An amniocentesis are performed, which reveals that the
fetus has AB blood. What is the chance that this fetus has nail-patella syndrome.
10.Elliptocytosis is hereditary trait in humans that is characterized by the presence of
oval-shaped red blood cells, rather than the normal biconcave-shaped cell. The gene
for elliptocytosis is dominant to its normal allele and is located on chromosome 1,
about 20 map units away from the Rh locus. The following pedigree shows the joint
inheritance of the gene for RH factor and elliptocytosis in three generation of a family.
The filled-in symbols represent individuals with elliptocytosis, while the presence or
absence of Rh factor is shown below each symbol. (a) Determine the genotypes of the
individuals in the pedigree. (b) What is the linkage phase (cis or trans) of individual II2? (c) Which individuals in generation III are recombinant types? (d) Calculate the
recombination frequency for the progeny in generation III. How does your estimated
map distance compare with the value given earlier in the problem?
Rh+

Rh-

Rh+

Rh+

Rh-

Rh- Rh- Rh+ Rh- Rh+ Rh- Rh+ Rh+

11.The recessive genes from hemophilia A and red-green color blindness in human are
located about 10 map units apart on the X chromosome. The ABO locus is on
chromosome 19. A normal man with type AB blood marries a normal , type AB woman
whose mother was colorblind and whose father was hemophilic. They have several
children. (a) What is the chance that their first child is color blind? (b) What is the
chance that their second child is a boy with type A blood who is both hemophilic and

color blind? (c) What is the chance that the third child will, like the parents, be normal
in color vision and clotting time and AB in blood type?
12.In corn, the genes for colored kernels (C) vs colorless (cc) and plump kernels (Sh) Vs
Shrunken (sh) are about 4 map units apart on chromosome 9. The dihybrid cross C
sh//c Sh X C sh//c Sh is made. (a) compute the genotypic ratio in the gametes
produced by each of the parents in the cross. (b) What percentage of the offspring will
be c sh//c sh in genotype? (c) What percentage of the offspring will have the colored,
shrunken phenotype? (d) What percentage of the offspring will be true-breeding?
13.In Drosophila, crossing over occurs in females but not in males. Suppose that the
dihybrid cross pr+ b//pr b+ X pr+ b//pr b+ is made, in which the genes for the
dominant red eyes (pr+) vs purple (pr) and fro the dominant gay body (b+) vs black (b)
are in the trans arrangement. The genes for these traits are located about 6 map units
apart on chromosome II. (a) Predict the phenotypic ratio among the progeny of this
cross. (b) Does the ratio you predicted indicate anything about how far apart the pr
and b loci are on the chromosome II? Explain.
14.The plants that are dihybrid for different pairs of gene loci are selfed. The frequency of
double-recessive offspring produced by each mating is given below. In each case,
determine whether or not the genes are linked. If the genes are linked, evaluate the
linkage phase (cis or trans) in the dihybrid parent, and calculate the map distance
between the genes involved.
Genotype of selfPollinated plant :
AaBb BbCc CcDd
Frequency of of doubleRecessive offspring: 1.00% 6.25% 9.00%
(aabb) (bbcc) (ccdd)
15. A testcross was made using the trihybrid PpRrSs as the dominant parent. The test
cross results revealed that the trihybrid produces the following gametes:
PRS
11 pRS
15
PRs
238 pRs 240
PrS
242 prS
230
Prs
11 prs
13
(a) Which loci are linked, and which assort independently? (b) Determine the
arrangement (cis or trans) of the linked genes in the trihybrid parent. (c) Calculate the
map distances between all linked loci.
16.In Drosophila, the three gene pairs for red-eyes (cn+) vs cinnabar (cn), normal bristle
number (rd+) vs reduced (rd), and the long wings (vg+) vs vestigial (vg) are known to
have their loci on chromosome II. Suppose that a three-point test cross yields the
following offspring:
Cinnabar, reduced, vestigial
406
Cinnabar, reduced, long
46
Cinnabar, normal, vestigial
28
Cinnabar, normal, long
3
Red, normal, long
438
Red, normal, vestigial
45
Red, reduced, long
33
Rd, reduced, vestigial
1
(a) Which are progeny classes of the parental type? Which are the single recombinant
types? Which are the double recombinant types? (b) Calculate the map distances

between the genes, and construct a linkage map of these loci. (c) Identify the most
probable origin of each of the recombinant classes, with regard to the number of
location of crossovers. (d) Determine the coefficient of coincidence for this set of loci.
17.In the seedling stage, a completely recessive corn plant was glossy (leaves have a
shiny appearance), virescent (poor in chlorophyll), and liguleless (lacking certain
appendages at the base of the leaves). This plant is crossed to a trihybrid that as a
seedling had dull leaves, normal chlorophyll content, and ligules. Of the many
seedlings produced in the next generation, a random sample of 1000 had the following
characteristics:
Dull, normal, with ligules
28
Dull, normal, liguleless
179
Dull, virescent, with ligules
69
Dull, virescent, liguleless
250
Glossy, normal, with ligules
198
Glossy, normal, liguleless
70
Glossy, virescent, with ligules
183
Glossy, virescent, liguleless
23
(a)Calculate the map distances between the genes, and construct a linkage map of
these loci (glossy = gl, virescent = v, and liguleless = lg). (b) Give the genotype of the
trihybrid parent, designating the proper gene arrangement on the pair of
chromosomes.
18. Assume the following linkage map:
a

10

6c

(a)Calculate the frequencies of the parental, single recombinant, and double

recombinant classes expected among the progeny of the three-point test cross
ABC//abc X abc//abc, assuming that crossing over occurs without interference. (b)
Repeat your calculations, but now assume interference, with a coefficient coincidence
of 0.5.
19. Three two-point testcrosses gave the following recombination frequencies:
Rp-q = 12%

Rq-r = 8%

Rp-r = 5%.

(a) Construct a linkage map af the p. q. r loci. Why are the distances not additive?
(b) Calculate the frequencies of the parental, single-recombinant, and doublerecombinant classes expected among the progeny of the three-point testcross PpQqRr
x ppqqrr, base on the values of Rp-q , Rq-r ,and Rp-r given above. (c) Determine the
coefficient of coincidence for this set of loci.
20. Consider the four-point testcross ABCD//abcd X abcd//abcd. (a) How many
recombinant classes can we possibly detect in the progeny of this cross? (b) Indentify
the most probable origin of each recombinant class, with regard to the number and
location of crossovers. (c) Suppose that data from previous two-point crosses indicate
that the members of each of the pairs of gene loci (a-b, b-c, and c-d) are two map units
apart. How many offspring must we examine from this four-point cross in order to
observe about eight individuals in the lowest frequency class?
21. You discover a recessive mutant gene in Drosophila that that is not reported in the
scientific literature, and you want to determine on which chromosome this gene is

located. Assume that you have access to two indicator strains of flies, one homozygous
for a gene on chromosome II, and another for a gene on chromosome III. Describe an
experimental breeding procedure you could use that would permit you to assign this
mutant gene to one of the four chromosomes in this species.
22. In a series of experiments with Neurospora, the following numbers of FDS and SDS asci
were observed for each gene pair. Determine the distance between the gene and its
centromere in each case.

NUMBER OF
TETRADS
FDS
SDS

GENE PAIR
(a) f+ vs. f (fluffy) growth

82

118

(b) c+ vs. c (colonial) growth 78

16

(c) r+ vs. r (ropy) growth

42

67

23. Neurospora of opposite mating types with the genotypes CD an cd are crossed.
Analysis of the 389 ordered tetrads produced from this cross yields the following data:
(1)

(2)

(3)

(4)

(5)

(6)

(7)

CD

Cd

CD

CD

CD

Cd

CD

CD

Cd

cD

Cd

Cd

cD

Cd

Cd

cD

Cd

cD

CD

Cd

Cd

cd

cD

cd

cd

cd

cD

cD

259

14

29

65

Determine whether or not the genes are linked. If they are linked, calculate the
distances between the genes. What are the gene-centromere distances?
24. Gene pairs c+, c (c = compact growth) and leu+. leu (leu + requirement for the amino
acid leucine) are observed in a mating of Neurospora srains c+ leu+ X c leu. The
following classes of ordered tetrads are obtained:

(1)

(2)

(3)

(4)

(5)

(6)

(7)

+ + + leu+

++

+ + + leuc

+ + leuc leuc

c leuc

++ leuc

c leuc
367

++

++ leu+

++

+c leu

++ leu+ leu

+ + leuc leuc leuc

4

11

50

+c

60

+
6

Determine linkage relationships and the map distances between the genes.
25. Unordered tetrad analysis in yeast gives the following data:

(1)

(2)

(3)

+ b

+ +

+ b

+ b

+ +

+ +

a +

a +

a +

363

234

Evaluate these data, and determine the distance between genes if they are linked.
26. Explain the origin of each of the tetrad classes in problem 23 through 25 by
diagramming the events of meiosis (assortment, crossing over) that were
responsible for their reproduction.
27. Unordered tetrad analysis of a cross between a+b+c+ and abc strains in yeast yields
the following data. Determine the map distances (genes are linked in the order
given).
(1)

(2)

(3)

(4)

(5)

+++

+++

+++

+++

+++

+++

+bc

++c

+b+

+bc

abc

a++

ab+

a+c

a+c

abc

abc

abc

abc

ab+

412

24

48

(6)

(7)

(8)

(9)

++c

++c

+bc

++c

+b+

+bc

+bc

++c

a++

a++

a++

ab+

abc

ab+

a++

ab+

28. Unordered tetrad analysis of a cross between d+e+f+ and def strains in yeast gives
the following data (genes are linked but not necessarily in the order given).
Determine the correct gene order and genetic distances.

(1)

(2)

(3)

(4)

(5)

+++

+++

+++

+++

+++

+++

d+f

++f

d++

d+f

def

+e+

de+

+ef

+ef

def

def

def

def

de+

299

42

90

10

11

(6)

(7)

(8)

(9)

(10)

++f

++f

d+f

++f

d++

d++

d+f

d+f

++f

d+f

+e+

+e+

+e+

de+

+e+

def

de+

+e+

de+

+ef

10

12

20

29. Suppose that in a study involving chromosome-mediated gene transfer, human

metaphase chromosomes are used to transfer HPRT mouse cells to the HPRT +
condition. Of the transformed HPRT + cells, 12 percent also received the human gene
that codes for glucose 6-phosphate dehydrogenase (G6PD). In a few cases, the

HPRT+ transformants received the PGK gene. Using these data on co-transfer
frequencies, construct a linkage map of the three loci.
30. Human cells are fused with mouse cells, and the resulting hybrid cells are placed
into HAT medium. After extensive chromosome loss, sex different cell lines are
selected and tested for the presence of human chromosomes as well as for the
presence of four human enzymes, one of which is thymidine kinase (TK), The results
are tabulated below. (The presence of a chromosome or enzyme is indicated by +,
the absence by -.).

CELL LINES
(a) (b) (c) (d) (e) (f)
Human Enzymes E1
+ + -

Human Enzymes E2
- + + -

+ +

Human Enzymes E3
+ + -

Human Enzymes E4
+ + + + + +
Human Chromo 1 + + -

Human Chromo 11+ + -

Human Chromo17+ + + + + +
Human Chromo X -

+ + -

+ +

(a). Identify the chromosome that carries the gene for each enzyme. (b) Which of the
enzyme (E1, E2, E3, or E4) is thymidine kinase (TK)? How can you tell? (c) The order
enzymes tested for were lactate dehydrogenase (LDH), 6-phosphogluconate
dehydrogenase (6PGD) and phosphoglycerate kinase (PGK).