Beruflich Dokumente
Kultur Dokumente
C H A P T E R
5
Quantitative
Genetics
(Inheritance of Multiple Genes)
Contents
64
Table 5.1.
Quantitative genetics
1. Characters of kind.
2. Discontinuous variation; distinct
phenotypic classes.
3. Single gene effects.
1.
2.
4.
3.
5.
Characters of degree.
Continuous variations; phenotypic measurements form a spectrum.
Polygenic control; effects of single genes
too slight to be detected.
Concerned with population of organisms
consisting of all possible kinds of matings.
Statistical analyses give estimates of
population parameters such as the mean
and standard deviation.
HISTORICAL
In 1760, Joseph Kolreuter inadvertently reported first case of continuous variation due to
quantitative trait. He crossed the tall and dwarf varieties of tobacco, Nicotiana. The F1 plants were
intermediate in size between the two parent varieties. The F2 progeny showed a continuous gradation
from the size of the dwarf to that of the tall parent. Since the basic principles of genetics were yet not
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65
EXAMPLES OF QUANTITATIVE
INHERITANCE
1. Kernel Colour in Wheat
A whole grain or seed of a cereal plant such as
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66
corn, wheat, barley, etc., is called kernel. Kernel colour in wheat is a quantitative trait and its
inheritance was studied by Swedish geneticist H. Nilsson Ehle for the first time in 1908. When he
crossed a certain red strain to a white strain, he observed that the F1 was all light red and that
approximately 1/16 of the F2 was as extreme as the parents, i.e., 1/16 was white and 1/16 was red. He
interpreted these results in terms of two genes, each with a pair of alleles exhibiting cumulative effect
(Fig. 5.1).
P:
Red kernel
White kernel
R1 R1 R2 R2
r1 r1 r2 r2
F1 :
Light red
R1 r1 R2 r2
F2 : Summary of checker board derived results, i.e., F2 genotypic and phenotypic ratios:
Genotype
Genotypic
ratio
R1 R1 R2 R2
R1 R1 R2 r2
R1 r1 R2 R2
2
2
R1 r1 R2 r2
R1 R1 r2 r2
r1 r1 R2 R2
4
1
1
R1 r1 r2 r2
r1 r1 R2 r2
2
2
r1 r1 r2 r2
Fig. 5.1.
Number of
contributing alleles
Phenotype
Phenotypic
ratio
Red
Medium red
Light red
White
Coloured
(15/16)
Colourless
(1/16)
Results of a cross between two varieties of wheat having red kernel and white kernel
showing cumulative effect of alleles.
Each of the contributing alleles R1 or R2 adds some red to the phenotype of kernel colour, so that
the genotypes of whites contain neither of these alleles and a red genotype contains only R1 and R2
alleles. These results are plotted as histograms in Fig. 5.2. Here five phenotypic classes are obtained
in F2; each dose of a contributing allele for pigment production increases depth of colour. At this stage
one point should be clear that in case there were two genes involved, there would be obtained 15 : 1
ratio (15 coloured : 1) (see Fig. 5.1).
Later on, when certain other strains of
wheat with dark red kernels were crossed with
P
1
whites exhibited an F1 phenotype intermediate
between the two parental types, but only
Dark red
1/64 of the F2 are white. In this case the F1 is
Red
probably segregating for three pairs of genes
Medium red and only the genotype r1r1 r2r2 r3r3 produce
F1
white. There are seven classes of phenotypes in
Light red
a ratio of 1 : 6 : 15 : 20 : 15 : 6 : 1.
Above described ratios, i.e., 1 : 4 : 6 : 4 :
White
1 and 1 : 6 : 15 : 20 : 15 : 6 : 1, can be easily
obtained by the expansion of binomial equation
F
2
(1/2+1/2)n , where n is the number of alleles. In
Fig. 5.2. Wheat colour as an example
case of 2 genes, n=4, while in case of 3 genes
of a quantitative trait (after
n=6. This expansion can be obtained by the use
Stansfield, 1969).
of Pascals triangle (see Table 5-2).
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E.M. East (1913) extended the polygenic hypothesis to several cases in plants. For instance, in
case of maize it was demonstrated that the ear size is controlled by multiple factors. Similarly, flower
size in tobacco had the same pattern of inheritance. In these cases, the number of genes controlling the
character were many and usually more than two or three.
Table 5.2.
Expansion of binomial (1/2+1/2)n using different values of n (number of alleles) with the
help of Pascals triangle.
Expansion of binomial (1/2+1/2)n
Number of alleles
1
2
3
4
5
6
7
8
1:1
1:2:1
1:3:3:1
1:4:6:4:1
1 : 5 : 10 : 10 : 5 : 1
1 : 6 : 15 : 20 : 15 : 6 : 1
1 : 7 : 21 : 35 : 35 : 21 : 7 : 1
1 : 8 : 28 : 56 : 70 : 56 : 28 : 8 : 1
AABB
Negro
aabb
White
AB
ab
P1 gametes :
F1 :
Intercross :
AaBb
Mulattoes
Intermediate skin colour
Aa Bb
Mulattoes
AaBb
Mulattoes
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&
AB
Ab
AABB
Like negro
AB
AABb
AABb
AaBB
Darker than mulattoes Darker than mulattoes
ab
AaBb
Like mulattoes
AAbb
AaBb
Like mulattoes
Like mulattoes
AaBb
aB Darker than mulattoes
AaBb
Like mulattoes
aaBB
Like mulattoes
aaBb
Lighter than mulattoes
ab
AaBb
Like mulattoes
Negro
colour
(1/16)
F2 :
Fig. 5.3.
aB
Aabb
Aabb
aaBb
Lighter than mulattoes Lighter than mulattoes
Colour between
mulattoes and negro
(4/16)
Colour of
mulattoes
(6/16)
Colour between
mulattoes and white
(4/16)
aabb
Like white
White skin
colour
(1/16)
Table 5.3.
Genotypes
Genotypic Frequency
Ratio
Black (Negro)
Dark
AABB
Aa BB,
AA Bb
1
2
2
1
4
Intermediate (Mulatto)
Aa Bb
aa BB
AA bb
4
1
1
Light
Aa bb
aa Bb
aa bb
2
2
1
White
These results are clearly showing that A and B genes produce about the same amount of
darkening of the skin and, therefore, the increase or decrease of A and B genes cause variable
phenotypes in F2 in the ratio of 1 Negro : 4 dark : 6 intermediate : 4 light : 1 white.
Other examples of quantitative traits of human beings include height, intelligence (I.Q.), hair
colour (except for red versus non-red) and eye colour.
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Table 5.4.
69
Number of contributing alleles for each type of eye colour of human beings (Source :
Burns and Bottino, 1989).
Eye colour
0
1
2
3
4
5
6
7
8
Light blue
Medium blue
Dark blue
Grey
Green
Hazel
Light brown
Medium brown
Dark brown
TRANSGRESSIVE VARIATION
In some types of quantitative inheritances, sometimes, F2 offsprings do not display continuous
variation but some individuals exhibit great degree of variability and do not resemble with their either
parent or even remote ancestors in that quantitative trait. Such a case in which the extremes of F2 exceed
those of the parent is called transgressive variation.
Example. Punnett and Bailey (1914, 1923) have reported first case of transgressive variation
from a cross in between a large Golden Hamburg chicken with the smaller Sebright Bantam variety of
chicken. The F1 was intermediate in size between the parents and fairly uniform, but mean size of the
F2 was about the same as that of the F1, but the variability of the F2 was so great that a few individuals
were found to exceed the size of either parental type (showing transgressive variation). In this case, four
gene loci are thought to cause the transgressive variation as follows :
P:
aa BB CC DD
Large Golden
Hamburg chicken
(6 contributing
alleles)
AA bb cc dd
Small Sebright
Bantam chicken
(2 contributing allele)
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F1 :
Aa Bb Cc Dd
Intermediate sized hybrid
(4 contributing allele)
F2 : Some genotypes could segregate out in the F2 with phenotypic values which exceed that
of the parents. For example :
AA BB CC DD
8 Contributing alleles
Larger than Golden Hamburg
Aa BB CC DD
7
,,
,,
Aa bb cc dd
aa bb cc dd
1
0
,,
,,
No contributing allele
(physiological minimum)
}
}
2.
3.
4.
(a)
(b)
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7.
71
Show, by means of appropriate genotypes, how parents may have children taller than themselves.
Skin colour in man is controlled by additive genes : If both mother and father have intermediate skin
colour, can you expect children (i) with lighter skin, (ii) with darker skin ? Can you expect children
with darker skin, if both parents have light skin ?
Two homozygous varieties of Nicotiana longiflora have mean corolla lengths of 40.5 mm and 93.3
mm. The average of the F1 hybrids form these two varieties was of intermediate length. Among 444
F2 plants, none was found to have flowers either as long or as short as the average of the parental
varieties. Estimate the minimal number of pairs of alleles segregating from the F1.
ANSWERS TO PROBLEMS
4.
5.
7.
All traits showing practically continuous variation are likely to be due to polygenes. Here, the traits
of intelligence, height, skin colour, and eye colour probably involve polygenes.
Any parental genotypes that can produce at least some progeny with a greater number of contributing
alleles than they themselves have are possible, for example, Aa Bb Cc Dd Aa Bb Cc Dd, Aa Bb
Cc Dd aa bb Cc Dd, etc.
If four pairs of alleles were segregating from the F1, we expect (1/4)4 = 1/256 of the F2 to be as extreme
as one of the other parental average. Likewise, if five pairs of alleles were segregating, we expect
(1/4)5 = 1/1024 of the F2 to be as extreme as one parent or the other. Since none of the 444 F2 plants
had flowers this extreme, more than four loci (minimum of five loci) are probably segregating from
the F1.