4.

INHERITANCE OF TRAITS (CHARACTERS)

4.1 Quantitative and Qualitative Traits:

4.1.1 Introduction:
(a) ● Trait: Is a certain morphological, anatomical or physiological feature of an organism. Examples are height,
weight, shape, colour, scent, texture, yielding capacity, resistance, days to maturity, etc.
● Controlled by a gene or genes, most of them are influenced by the environmental factors.
(b) Genetic inheritance is broadly divided into quantitative and qualitative inherited traits.
● Attributes of Quantitative and Qualitative Traits:
Qualitative traits Quantitative traits (or Metric traits)
(i) Phenotype: clearly separated into discrete/distinct - differentiated by measurement; characterized
classes that do not overlap, as illustrated below: by a continuous range of the expression of the
phenotype, as illustrated below:

Aa AA+Aa

AA aa aa

No dominance With dominance Unit sizes

(1: 2:1) (3:1)
(ii) Least affected by environmental factors, e.g.
. flower colour, seed shape etc.
(iii) Controlled by one or few major genes with
large, easily detectable effects, i.e. oligogenes
- Easily affected by environmental factors, e.g. yield,
height, days to flowering, etc.
- Governed by several minor genes, each contributing
a small effect, i.e. polygenes.

(Read - Multiple Factor Hypothesis)

4.1.2 Population and Quantitative Genetics:

(a) Concepts:
(i) Population genetics and quantitative genetics:-
● Both deal with the genetic basis of phenotypic variation among the individuals in a population.
● Population genetics - focuses on frequencies of alleles and genotypes.
● Quantitative genetics – deals with the ways in which individual variation in genotype and environment
contribute to variance in phenotype.
(ii) Gene pool: It is a collection of all the alleles of all the genes found within a freely interbreeding population of
an organism, e.g. a plant species.
(iii) Gene Frequency: Also referred to as Allele Frequency. It is the fraction of the genes in the gene pool that is
composed of an allele in question, or the proportion of an allele (A or a) in the population.
● Genotypic frequency: Proportion of a genotype, AA, Aa or aa, in the population.
(b) Factors that Affect Changes in Gene Frequency in a Population:
Evolution occurs when there are changes in the frequencies of alleles within a population. This happens
mainly through four main processes:
(1) Selection:
♦ It is the most important process for altering population gene frequencies by plant breeders.
♦ Achieved through,
 (i) Natural Selection – through environmental factors; where it converts differences in fitness into changes in
allele frequency in a population over successive generations.
(ii) Artificial Selection – implies directional selection by mankind (plant breeders); based on defined objectives
and procedures.
(2) Mutation: It may produce a new allele not present in the population, or may change the frequencies of
existing alleles.
(3) Genetic Drift:
♦ It is a random change in gene frequency due to sampling error – which is greater in smaller populations.
Eventually, the frequency of one allele becomes zero (0), and that of the other allele becomes one (1) – i.e.
fixed in the population.
♦ Breeding populations are generally small, hence genetic drift is expected.
(4) Gene Flow and Transfer:
♦ Gene flow: refers to the exchange of genes between populations – usually of the same species. This occurs
through,
(i) Migration – movement of individuals into or out of an existing population.
(ii) Hybridization – incorporation of a gene(s) through cross-pollination.
♦ Gene transfer: implies incorporation of a gene using biotechnological techniques, e.g. genetic engineering,
which facilitate direct gene transfer – even if the species are different.

● The Hardy-Weinberg Principle (Equilibrium, Law, Model):

► It is the fundamental law of population genetics; provides the basis of studying Mendelian Inheritance.
► According to this principle, the frequencies of alleles will remain constant in the absence of selection,
mutation, genetic drift and gene flow/transfer.
► The principle also concerns the effects of a single generation of random mating, which can be expressed
mathematically as follows:
♦ Consider a single locus with two alleles, A and a, with their frequencies denoted by p and q, respectively, i.e.
freq (A) = p, freq (a) = q.
♦ The proportions of genotypes resulting from the mating and their frequencies in F 2 segregating population will
be, AA = p2, aa = q2 and Aa = 2pq.
♦ The allele frequencies must add up to one (unity) as follows:
p + q = 1; since (p + q)2 = p2 + 2pq + q2, hence p2 + 2pq + q2 = 1 (or 100%).

(c) Calculation of Gene Frequency:

● The above Hardy-Weinberg Equilibrium assumes dominance at the locus.
● Assume the genotypic frequencies are D for AA, H for Aa and Q for aa. Since the population is diploid, there
will be 2N alleles in it. The genotype AA has two A alleles, hence the total number of A alleles in the
population is equal to 2D + H.
● The frequency (proportion) of A alleles, designated as p in the population is therefore obtained as follows: (2D
+ H)/2N = (D + ½H)/N = p.
Likewise, for a allele, designated as q, will be (2Q + H)/2N = (Q + ½H)/N =q.

♦NB: Illustration:
Genotype AA Aa aa
Genotypic freq. p2 2pq q2
● Examples:
(1) If N = 80, D =4 and H = 24, then p = (D + ½H)/N = (4 + 12)/80 = 0.2.
Since p +q = 1, q = 1 – p, hence q = 1 – 0.2 = 0.8.

(2) Calculating genotypic frequency for the next generation in a random mating in the above example:
AA = p2 = 0.22 = 0.04
Aa = 2pq = 2(0.2 x 0.8) = 0.32
aa = q2 = 0.82
= 0.64
Total = 1.00
(d) Gene action (effect): There are 4 types of gene action, (1) Additive gene action, (2) Dominance gene action,
(3) Overdominance gene action and (4) Epistatic gene action.
The first three occur within the locus, and the last one between loci, as explained below.

Aa Aa Aa Aa
● ● ● ● ● ● ●
6a 5 4 3 2 1 6b
AA mid-parent aa
value
(1) Additive gene action (no dominance): Each allele contributes one increment to the expression of a trait,
hence heterozygote = mid-parent value, i.e. Aa = 3.
(2) Dominance gene action (effect): Are the deviations from the additivity that make the progeny from a
cross resemble one parent more than the other, e.g.
♦ Partial dominance effects, Aa = 4.
♦ Complete dominance effects, Aa = 5.
(3) Overdominance gene action: Examplified by Aa = 6a and/or 6b, i.e. heterozygote exceeds either parent.
The combined effect of each allele at a locus exceeds the independent effect of the alleles. This is a case of
TRANSGRESSIVE SEGREGATION (VARIATION), which means segregation/variation outside the
range of parents.
(4) Epistatic gene action (effect): Epistasis = expression of genes at one locus is influenced by the genes at a
different locus; though the genes are inherited independently, but interact to form one effect.
● Factors that affect gene action: (1) type of genetic material, (2) mode of pollination, (3) mode of inheritance
and (4) presence of gene linkage.

(e) Genetic components of continuous variation:

(i) Recall VP = VG + VE. Further, recall that the genotype and the environment are not independent of each other,
they do interact (G x E or GE); hence, VP = VG + VE + VGE.
● VG can be partitioned into, VG = VA + VD + VI, where ,
VA = variance due to additive gene action.
VD = variance due to dominance gene action.
VI = variance due to genetic interaction among loci.
● Thus VP = VA + VD + VI + VE + VGE.
 ● VA is the most prominent; contribution of VD and VI is very small – depending on the heterozygosity of the
individual.

4.2 Heritability:
(a) Concept: ● It refers to that proportion of the observed phenotypic variation for a given trait in a population, that
is due to genetic control (i.e. heritable).
● Designated H or h2.
● Formula, H or h2 = VG/VP, where, VG = genetic variance and VP = phenotypic variance.
● It is a ratio measured on a scale of 0 (zero) to 1 (one), where,
0 = all variations are due to environmental effects,
1 = all variations are due to genetic differences.
(b) Types of heritability:
(i) Broad sense heritability, h2b or Hb.= VG/VP; which estimates h2 on the basis of all genetic effects.
(ii) Narrow sense heritability, h2ns or Hns = VA/VP, this estimates h2 on the basis of additive gene effects.
(c) Estimation/calculation of h2: To be done after covering topic 5.

(d) Importance of h2 in plant breeding:

(i) To determine the most effective selection strategy/method in a plant breeding program. Example,
for low h2, use more sensitive selection methods, whereas for high h 2 use any selection method.
(ii) Used in predicting response to selection (gain from selection).
N.B. Heritability will not be the same in populations of similar plant species in different locations, due to differences
in environmental factors.

4.3 Correlation of traits:

(a) Concept: It refers to the relationship or association between traits. Example, a breeder would like to know the
relationship, say between yield and plant height, protein content and oil content, protein content and seed size,
etc., i.e. influence of one trait over the other.
(b) Measured by (1) correlation coefficient (2) path coefficient and (3) discriminant function analyses.
● Correlation coefficient analysis: three types, (i) simple linear correlation, (ii) partial correlation and (iii)
multiple correlation.

● Formula for simple linear correlation (SLC) is,

∑ 𝒙 .∑ 𝒚
𝒓 𝒄𝒐𝒗 𝒙.𝒚 𝒓 ∑ 𝒙𝒚
(±) = or (±) = 𝒏
(∑ 𝒙)𝟐 (∑ 𝒚)𝟐
𝑺𝟐𝒙 .𝑺𝟐𝒚 ∑𝒙 𝟐 .∑𝒙 𝟐
𝒏 𝒏
Where, cov.x.y – covariance for traits x and y.
𝑺𝟐𝒙 and 𝑺𝟐𝒚 – variances for x and y traits.
● r ranges from +1 to 0 to -1.
● If the correlation is –ve, then an increase in one results into a decrease in the other, e.g. in soybean,
↑ protein content → ↓ oil content.

(c) Useful in indirect selection of traits.

(d) Causes of correlations:

(i) Pleiotropy: a condition where a single major gene affects two or more traits, i.e. extent to which two (or >)
traits are influenced by a gene.
(ii) Gene linkage: a particular set of characters may be related because genes are linked.
 (iii) When traits share the same resources for development, generated by a given set of genes.

5. BASIC STATISTICAL CONCEPTS IN PLANT BREEDING

5.1 Concepts: What do you understand by the following terms as used in statistics? Statistics, parameter,
variable, population, sample.
5.2 Measurements:
N.B.: Take note of symbols of measures.
5.2.1 Measures of Central Tendency: Mean, median and mode. They do not indicate anything about the spread
of the values.
5.2.2 Measures of Dispersion/Variability: Concerns the degree to which values of a data set differ from their
computed mean or differ from each other.
(a) Range: Is the difference between the highest and the lowest values of a sample. It concentrates entirely on the
extreme ends.
Example, (i) 12.3, 13.2, 12.0, 12.9, 13.8; range from 12.0 to 13.8 and a mean of 12.84
(ii) 17.4, 6.9, 6.0, 19.8, 14.1; range from 6.0 to 19.8 and a mean of 12.84.
▪ Which one has a greater variability?
(b) Variance (𝝈², 𝒔²): Also called mean square.
♦ Expressed in the units of the observation, squared, e.g. cm 2,kg2, etc.
♦ Magnitude of variance reflects the scatter in the data; a larger variance reflects larger deviations.
(∑ 𝒙)²
∑ 𝒙² ∑(𝒙 𝒙)²
♦ Formula: S² = 𝒏
or S² =
𝒏 𝟏 𝒏 𝟏

▪ N.B.: (n-1) = degree of freedom (df) → the concept is that in any given sample set, there is one value that is
not independent. The use of df gives an unbiased estimation of the variance.
♦ Example: Data on plant height (cm): 25, 32 29, 26, 31, 27, 30, 28.
(∑ 𝒙)²
∑ 𝒙² (∑ 𝒙)²
▪ S² = 𝒏
, Where n = 8, ∑𝒙𝟐 = 𝟔𝟓𝟒𝟎, = 𝟔𝟒𝟗𝟖
𝒏 𝟏 𝒏

𝟔𝟓𝟒𝟎 𝟔𝟒𝟗𝟖 𝟒𝟐
∴ S² = = = 6cm2
𝟖 𝟏 𝟕

(c) Standard deviation ( 𝝈, 𝑺 𝒐𝒓 𝑺𝑫/𝑺𝒅):

♦ It is the square root of the variance. Expressed in the same units of measurement as the observation, hence
preferred to the variance as a descriptive measure. S 2 and Sd are complementary.
♦ Measures variability that indicates by how much the value in a distribution deviates from the mean.
 (∑ 𝒙)²
∑ 𝒙²
♦ Formula: S or Sd = 𝑺², i.e. S = 𝒏
𝒏 𝟏

▪ From the above data, S = √𝟔 = 2.45cm

(d) Standard error (SE or 𝑺𝒙 ):

♦ Also called standard deviation of the mean or standard error of the mean. It applies to the means as opposed
to the Sd which applies to the observations.
♦ It is the measure of uncontrolled variation present in a sample.

𝑺 𝑺𝟐
♦ Formula: SE = or SE =
√𝒏 𝒏

𝟐.𝟒𝟓
▪ From the above data, SE = = 𝟎. 𝟖𝟕
√𝟖
(e) Coefficient of Variation (CV):
♦ It is a measure of the relative variability of given populations or traits in a given population.
♦ It is also an index of the reliability of the experiment, as it indicates the degree of precision with which data
was collected. Higher CV implies that data was unreliable. Generally CV of ≤ 20% is acceptable,
depending on the experiment; the lower the better.
𝒔 𝟐.𝟒𝟓𝒄𝒎
♦ Formula: CV = x 100; from the above data, CV = x 100 = 8.6%.
𝒙 𝟐𝟖.𝟓𝟎𝒄𝒎

5.3 Experimental Designs, Data Collection, Analysis and Interpretation:

5.3.1 Introduction:
(a) Why analyse data? It is done to get/determine the difference among the treatments.
(b) Methods of analysis: Differ with; (i) Number of treatments (ii) Experimental design used.
(c) Experimental designs:
(i) Types – CRD, RCBD, Latin square, etc.
(ii) Terms: (1) Experimental unit/plot (2) Replication (3) Treatment (4) Randomization
(5) Experimental error (6) Local control.

5.3.2 Data Analysis and Interpretation:

(a) Data collection: (i) Sampling (ii) tabulation, e.g. Root length in chickpea (cm).

Treatment Replications Treatment Treatment

totals means
I II III
A 18.22 14.33 16.08 48.63 16.45
B 9.61 7.92 11.08 28.61 9.54
C 16.05 15.72 13.16 44.93 14.00
D 10.00 12.82 11.33 34.15 11.38
E 7.94 6.57 10.06 24.57 8.19
F 19.65 17.00 18.03 54.68 18.23
Repl total 81.47 74.36 79.74
 Grand total (GT) 235.57

(b) Data Analysis and Interpretation:

♦ Steps:
(1) Calculate correction factor (C.F. or C)
(𝐆𝐓)𝟐 (𝟐𝟑𝟓.𝟓𝟕)𝟐
- C.F. = (𝒓𝒙𝒕)
⟹ (𝟑𝒙𝟔)
= 3082.96

(2) Calculate Replicate Sum of Squares (RSS)

𝑻𝟐𝑹𝟏 𝑻𝟐𝑹𝟐 ...𝑻𝟐𝑹𝒏
- RSS= − 𝑪. 𝑭 Where TR1,TR2, etc = totals of replicates
𝒕
t = # of treatments
𝟖𝟏.𝟒𝟕² 𝟕𝟒.𝟑𝟔² 𝟕𝟗.𝟕𝟒²
 RSS = − 𝟑𝟎𝟖𝟐. 𝟗𝟔 = 𝟒. 𝟓𝟖
𝟔

(3) Calculate Treatment Sum of Squares (TtSS)

𝑻𝟐𝒕𝟏 𝑻𝟐𝒕𝟐 ...𝑻𝟐𝒕𝒏
- TtSS = − 𝑪. 𝑭, Where Tt1, Tt2, etc = treatment totals
𝒓
r = # of replications
𝟒𝟖.𝟔𝟑𝟐 𝟐𝟖.𝟔𝟏𝟐 ⋯ 𝟓𝟒.𝟔𝟖𝟐
 TtSS = − 𝟑𝟎𝟖𝟐. 𝟗𝟔 = 𝟐𝟑𝟕. 𝟔𝟖
𝟑

(4) Calculate Total Sum of Squares (TSS)

- TSS = 𝒙𝟐𝟏 + 𝒙𝟐𝟐 + 𝒙𝟐𝟑 . . 𝒙𝟐𝒏 − 𝑪. 𝑭, where x1, x2, etc = are the mean observations in each experimental unit.
 TSS = (𝟏𝟖. 𝟐𝟐² + 𝟏𝟒. 𝟑𝟑²+. . . 𝟏𝟖. 𝟎𝟑²) − 𝟑𝟎𝟖𝟐. 𝟗𝟔 = 𝟐𝟔𝟗. 𝟎𝟏
(5) Calculate Error Sum of Squares (ESS)
- ESS = TSS-RSS-TtSS
=> ESS = 269.01 - 4.58 - 237.68 = 26.75

(6) Analysis of Variance (ANOVA) table

Source of D.f. SS M.S Fcal Fcrit/tab

variation (n-1) (= Variance) 5%(0.05) 1%(0.01)
Replication (r-1) RSS 𝐑𝐒𝐒
= 𝐑𝐌𝐒
𝐝.𝐟
3-1= 2 = 4.58 𝟒.𝟓𝟖
𝟐.𝟐𝟗
𝟐
Treatment (t-1) TtSS 𝐓𝐭𝐒𝐒
= 𝐓𝐭𝐌𝐒
𝐓𝐭𝐌𝐒 3.33 5.64
𝐝.𝐟 𝐄𝐌𝐒
6-1 =5 = 237.68 𝟐𝟑𝟕.𝟔𝟖 𝟒𝟕.𝟓𝟒
𝟓 𝟐.𝟔𝟖
= 𝟒𝟕. 𝟓𝟒 = 𝟏𝟕. 𝟕𝟒
Error (r-1)(t-1) ESS 𝐄𝐒𝐒
= 𝐄𝐌𝐒 - - -
𝐝.𝐟
2 ˟ 5= 10 = 26.75 𝟐𝟔.𝟕𝟓
𝟏𝟎
= 𝟐. 𝟔𝟖
Total (rt)-1 TSS - - - -
(3˟6) -1= 17 = 269.01

(7) Interpretation of the ANOVA Results: Done from the comparison of Fcal and Fcrit/tab.
 You will declare that there is significant difference among treatments, IF Fcal > Fcrit/tab and vice versa, at say
5% (0.05) or 1% (0.01) level of probability or significance.
-Above case: There is significant difference among treatments at both 5% and 1% levels of probability.

(8) Mean Separation:

- Done only if there is significant difference among treatments.
- Methods:
1. Least significant difference( LSD) test
2. Duncan’s Multiple Range Test (DMRT)
3. Orthogonal Coefficients
4. Etc

Use of LSD:
𝟐𝑺²
(i) 𝑳𝑺𝑫∝ = 𝒕∝,𝒆𝒅𝒇 × , Where ∝ = Level of probability, edf = error degrees of freedom, t = t-test (read
𝒓
in the t-test table), S² = Variance = EMS, r= # of replications.
𝟐×𝟐.𝟔𝟖
(ii) 𝐋𝐒𝐃𝐨.𝐨𝟓 = 𝐭 𝟎.𝟎𝟓,𝟏𝟎 × = 𝟏. 𝟑𝟑𝟔𝟔𝟔𝟐𝟓
𝟑

− 𝒕𝟎.𝟎𝟓,𝟏𝟎 → Read from t table values; = 2.228; Therefore, LSD = 2.228 ×1.3366625 ≅ 2.98

(iii) Two means will be declared significantly different if their difference is greater than the LSD value-and vice
versa.

(iv)Presentation of Results: Arrange from the highest to the lowest mean, i.e.

Treatment: F A C D B E
Mean: 18.23a 16.45ab 14.00bc 11.38cd 9.54de 8.19e

*Depending on the trait and/or objective, treatments F and A are superior to the rest and vice versa, i.e.
treatments B and E is superior to the rest of the treatments.

CALCULATIONS OF HERITABILITY

A) Broad Sense Heritability (h2b):

(1) Using ANOVA table:

𝑽
♦ h2b = 𝑮 ; Recall, VP = VG + VE
𝑽𝑷

𝑻𝒕𝑴𝑺 𝑬𝑴𝑺
♦ VG = ; 𝒓 = # of replications.
𝒓

♦ VE = EMS

♦ Example: Calculate h2b using the above ANOVA table.

♦ Implication: If h2 is high, it means that the genotype plays a more important role than the environmental effect
in determining the phenotype, i.e., a trait with high h 2 will respond to selection more than the trait with low h 2.

(2) Using variances of parents 1 and 2 (P1 & P2), F1 and F2 populations:
♦ Estimate variances of P1, P2, F1 & F2 separately.

♦ P1, P2 & F1 are non-segregating populations; hence any variation will be due to the environmental effects
𝑽𝑷𝟏 𝑽𝑷𝟐 𝑽𝑭𝟏
only. Therefore, VP1, VP2 & VF1 will give an estimation of VE, e.g. VE =
𝟑
OR
𝑽𝑷𝟏 𝑽𝑷𝟐
VE = OR VE = VF1.
𝟐

♦ F2 is a segregating generation/population; hence variation will be due to both genetic and environmental effects.
Therefore, VF2 = VG + VE. This implies that VF2 = VP, and also VG = VF2 – VE.

𝑽𝑮 𝑽𝑭𝟐 𝑽𝑬
♦ Finally, h2b = or h2b = .
𝑽𝑷 𝑽𝑭𝟐

♦ Example: Past papers.

B) Narrow Sense Heritability (h2ns):

Using variances of F2 and backcrosses 1 & 2 (BC1 & BC2) populations:

𝟐𝑽𝑭𝟐 𝑽𝑩𝑪𝟏 𝑽𝑩𝑪𝟐

♦ h2ns = , where 𝑽𝑩𝑪𝟏 𝒂𝒏𝒅 𝑽𝑩𝑪𝟐 are the variances of backcrosses 1 & 2.
𝑽𝑭𝟐

Example: