Variance

Sirjan Bastola
2019-06-18
 Complex traits
- Traits influenced by many factors: multiple genes, interaction
between alleles of different genes, variations in environment
interaction between genes and environment, and chance events
- e.g. height of adult human

 Quantitative traits
- Many complex traits are quantitative traits
- Quantitative approach of measurement  Phenotype can be
measured over a range of numbers called phenotypic values or trait values
- Show a roughly bell-shaped normal distribution of phenotypic values in
population (apparently continuous variation or distribution)
- e.g. body size, height, milk yield, enzyme activity, reproductive ability
 Quantitative traits are most often controlled by two/ more genes
that provide additive component to the phenotype that can be
quantified.
 Continuous variation characterizes the inheritance of quantitative traits
 Inheritance of quantitative traits= Quantitative/ Polygenic inheritance
Quantitative traits show normal
distribution (continuous variation)

Genetic and environmental factors

influence quantitative traits
 Multiple genes influence quantitative traits
(Multiple factor Hypothesis)

Figure. Inheritance of grain color in wheat. Three

independently assorting genes (A, B and C) are assumed
to control grain color. Each gene has two alleles. The alleles
that contribute additively to pigmentation are represented
by uppercase letters.
Figure. How multiple-factor hypothesis accounts for 1:4:6:4:1
phenotypic ratio of grain color when all alleles designated by
an uppercase letters are additive and contribute an equal
amount of pigment to the phenotype.
Figure. Histogram showing the relative frequency of individuals
expressing various height phenotypes derived from Kolreuter’s
cross between dwarf and tall tobacco plants carried to F2
generation.
 Statistics of Quantitative Genetics
 Biometry
- Quantitative, statistics- based approach to study the
inheritance of traits exhibiting continuous variation
 Frequency distribution
1) First step in study of quantitative traits is to select sample
2) Collect data and present in graph as frequency distribution
3) Frequency distribution = Different types of variation within
each sample

Figure. Normal frequency distribution,

characterized by a bell-shaped curve
 Mean
- Mean represents the clustering around central Value (Central tendency).
- Mean Provides a descriptive summary of sample but does not tell about the
range.
- Modal class is the class with most observation.

Figure. Two normal frequency distributions with

same mean but different amounts of variation
 Variance (Sample variance)
 Data in frequency distribution could be dispersed or clustered
 It is a statistic to measure the (degree of) spread of data in frequency distribution
i.e. divergence of values from mean
 Data that are widely dispersed produces a large value for variance
 Data that are tightly clustered produces a small value for variance
 It is used as an estimate of variation present in an infinitely large population
 2 features of variance
1) Measures the dispersion of date around mean
2) Variance is always positive Point of inflection

 Standard deviation
 It is the square root of variance.

Figure. A normal frequency distribution showing

the percentage of measurements within 1, 2 and 3
standard deviations of the mean.
 Standard error of mean
 It is a measure of accuracy of sample.
 It measures variation of sample means in
replication of experiments.
 It estimates how much the means of similar
samples drawn from same population might vary.

 2 Key points
 Mean class point to the center in frequency distribution.
 Variance and standard deviation are statistics that indicate the extent
to which data are scattered around the mean in frequency distribution.
 Multiple Factor (Gene) Hypothesis
Figure. Quantitative phenotypes and the deviations of individual
measurements from the population mean. Each individual’s
deviation is hypothesized to consist of a deviation due to its
genotype (g) and a deviation due to its environment (e).
 R.A Fisher proposed this hypothesis.
 Key idea- Certain traits are controlled by
multiple factors in the genotype and
environment.
 He proposed a particular value of a
quantitative trait is result of combined
influence of genetic and environmental
factors.
 He represented the effects of these factors
as deviations from overall population mean.
T=µ+g+e
T- Phenotypic value of a quantitative trait
µ- population mean
g- deviation from mean due to genetic factor
e- deviation from mean due to
environmental factor
(g and e vary with individuals)
 These traits are quantitative traits and many
genes (= polygenes) contribute to phenotype of
such traits in a cumulative/ quantitative way.
So, also known as polygenic traits.
 Multiple Factor (Gene) Hypothesis- Key points
(Also Additive gene action)

1) Character that exhibit continuous variation can be quantified (measure, weigh,

count).
2) 2 or more pairs of genes located throughout the genome account for the hereditary
influence in an additive way.
Many genes involved=Polygenic inheritance
3) Each gene locus may be occupied by
a) additive allele  contributes a constant amount to phenotype
b) non-additive allele  doesn’t contribute quantitatively to the phenotype

Additive alleles are the basis of continuous variation

4) Total effect of each additive allele ~ To all other additive alleles at other gene sites.
5) Together, genes controlling a single character produces substantial phenotypic
variation.
6) Analysis of polygenic traits requires the study of large number of progenies from a
population of organisms.
 Phenotypic Variance

Number of cows

Milk yield
Finding mean
Let xl = the milk yield of the cow l in a population of
N cows. The mean of milk yield, for the population
is defined as

Finding variance
The variance Vp of milk yield for the population
is defined as

 Variance is a statistical measure of the amount

of variation in a population.
 Partitioning Phenotypic Variance
1) Genetic variance (VG ) 2) Environmental variance (VE )
Genetically different cows reared in same climate Genetically identical cows reared in variable climate

Number of cows
Number of cows

Milk yield Milk yield

3) Phenotypic variance (VP = VG + VE )
Genetically different cows reared in variable climate
Number of cows
Figure: Genetic and environmental components of
phenotypic variance. 1) The phenotypic variance of
genetically different cows reared in same climate is
all due to the genetic variance VG . 2) The phenotypic

variance of genetically identical cows reared in variable

climate is all due to environmental variance VE. .
3) For natural population reared in diverse natural
environment, the total phenotypic VP is the sum of the
genetic and environmental variance components
(VG + VE )
 Genetic variance can be separated
Milk yield
from environmental variance.
 A
Breed

B
Breed
 The third component of phenotypic variation
Nutrition is genotype-by-environment interaction
variance (VG x E)
 We can now summarize all the components of
total phenotypic variance VP using the following
equation:
VP = VG + V E + VG x E
Breed
A  In other words, total phenotypic variance can be
subdivided into
B
ed - genotypic variance
Bre
- environmental variance, and
Nutrition - genotype-by-environment interaction variance
Figure. Differences in yield between two different cow breeds
at different nutrition levels. (a) No genotype-by-environment,
or G x E, interaction: The breeds show genetic differences in
yield but respond equally to increasing nutrition. (b) G x E
interaction present: Breed A outyields B at low nutrition, but
B yields more than A at high nutrition levels.
 Growth of tropically
adapted genotype

Temperature

Environmental effect Genotype by Environment Effect

(Effect of temperature on growth rate) (GXE interaction)
 Khaire grown and trained
in Lamjung

Holstein
Running velocity

Milk production
Gore grown and trained
in Rupandehi Haryana

Temperature Temperature
Environment effect Genotype by Environment effect

 For Genotype by Environment effect (G X E), there must be

two genotypes in two environment
 Phenotypic Variance

 VP = VG + VE + VGE
VP - (Total) phenotypic variance/ Variance of quantitative trait
VG - Genetic variance (measures the effects of genetic differences among individuals)
VE - Environmental variance (measures the effects of environmental differences among
individuals)
VGE - Variance resulting from the interaction of the genetics and environment
Since VGE is usually negligible, VP = VG + VE

 VG = VA + VD + VI
VA - Additive variance (variance due to additive genetic effects, results from average effect of
additive components of genes)
VD - Dominant variance (variance due to dominance effects, results when the phenotypic
expression of heterozygous is not precisely intermediate between 2 homozygotes)
VI - Interactive variance or Epistatic variance (variance due to non-additive interaction between
alleles at different loci or deviation from the additive components that occur when two/ more
loci behave epistatically)
 VG = VA + VD + VI  V =V +V
E TE PE

Additive Non-additive Temporary Permanent

variance variance environment environment
variance variance
What are the differences?