Agro 222 June 2021

6. SELECTION

6.1. Selection:
Concept: Process of isolating desirable/suitable genotypes from the undesirable ones, based on traits of
interest, in a population, e.g. of plants.
● Categorized into natural and artificial selection:
(a) Natural selection:
- Natural forces like climatic conditions, soil factors, competition for space and food, etc, eliminate the
poorly adapted variants and retain those that cope with these forces.
- No time frame, but notable changes are observed at some stage.
(b) Artificial selection:
- It is a purposeful process of selecting certain genotypes by mankind under a chosen or an identified set of
environments and/or objectives.
- Remarkable changes are noted at the end of the set time frame.
- Effectiveness depends on (i) extent of genetic variability and (ii) h 2 of character (trait)..
● Selection changes gene frequencies; thus creating new gene pools or plant ideotypes.
- Plant ideotype: A model/design of a desired plant phenotype, according to the purpose of the cultivar.
Various traits are selected for in developing a plant ideotype.

6.2. Selection Approaches:

●A plant breeder must first address the task of what traits are to be selected for; i.e. set the breeding
objectives. This will depict a model/design of a desired cultivar, i.e., a crop plant ideotype.
● Normally simultaneous selection is carried out, i.e. selection involving ≥2 traits (characters) that are
improved at the same time. This can be achieved by three approaches:
(i) Tandem selection: Each trait is selected singly in turn till it is improved, i.e. selection of one trait is
completed first and then followed by another.
(ii) Truncation (Independent culling) selection: Selection is done for all the traits at the same time, but
independently. This is done by rejecting all individuals that fail to meet a certain standard for each trait;
regardless of their values for any other trait they may possess.
(iii) Selection index: Selection is applied to all traits simultaneously, with appropriate weight given to each
(trait) based on its relative economic importance, its heritability and its genetic and phenotypic correlation
among these traits in question.
Formula: I = b1x1 + b2x2 + …+bnxn, where, I = selection index,
x1, x2, ..xn = phenotypic values of traits considered and b1, b2,...bn = corresponding weights, calculated
from a series of simultaneous equations involving the appropriate phenotypic and genotypic variances
and co-variances; which are to be estimated for each cycle of selection.

● Field Trials: (i) Refer to topic #5, section 5.3.

(ii) Types of field trials: (1) Preliminary yield trials, (2) Advanced yield trials, (3) Stability yield trials (4)
On-farm or adaptive trials (5) National performance trials.

6.3. Response to Selection:

(a) - The effect of selection is to change the relative frequency of the different alleles at a segregating locus,
hence the observed change in the phenotypic means of selected traits.
- Response to selection will be shown by an increase or a decrease in the mean (𝑿) of the selected trait.

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(b) Concept / definition:

● Response to selection: It is the difference in the phenotypic value (depicted by means, variances e.t.c.)
between the offspring of the selected parents and the initial (original) population from which parents were
selected from.
It is also referred to as genetic gain (∆𝐆) or genetic advance (∆𝐆).
♦Illustrations:
(i) Formula: R = 𝑿OF – 𝑿I, where R = response to selection, 𝑿OF = mean of offspring population and
𝑿I = mean of initial population.

(ii) Graphical:

Selected parents (e.g. 5% of the total population)

C0

𝑿I S 𝑿P

Advance/gain
C1 made
- Offsprings of the selected parents.
- Note the shift in the mean of C1 popln compared to that of C0.

𝑿OF

Where, 𝑿P = mean of selected parents.

S = Selection differential (𝑿P – 𝑿I).
C0 = Selection cycle 0 (zero).
C1 = Selection cycle 1 (one).

(c) Response to selection or genetic gain or genetic advance under selection is related to heritability by
the following equation/expression, R = h2.S, where h2 is the heritability of the trait under selection, and
S is the selection differential.

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(d) The linear response seen in early generations of selection is not maintained indefinitely. This can be
depicted graphically as follows:

I
F
C E G H
B D
A

0 10 20 30 40 50 60 70 80 90
Generations of selection
Key:
A = No selection applied, population mean fluctuates about a constant level.
B = Steady linear response; change of genes frequency influencing the trait.
C = Gene frequencies reach high level; some loci become fixed in homozygous state, hence the plateau.
D = Natural selection may cause a gradual decline.
E = Some event may occur which releases variability, hence a linear response phase.
ETC.

(e) Maximizing response to selection:

● Recall R = h2.S
● If both sides are divided by the phenotypic Standard deviation, S.D. (𝛔), then we shall end up proving
that, R = 𝛔.i.h2 or ∆𝐆 = 𝛔.i.h2 ,where,
(a) i = S/𝝈 (or standardized selection differential), it is a constant based on selection intensity
(selection pressure), i.e. the proportion (percentage) selected as parents from the population.
(b) 𝝈 = Standard deviation; it is dependent on genetic variation of the base population.
(c) h2 = heritability of trait under selection.
● Therefore R can be maximized by manipulating the three components:
▪ h2: It can be increased by minimizing environmental influences, e.g. by increased replications, blocking
in the field, etc.
▪ i: The smaller the proportion/percentage selected as parents the larger the value of i and vice versa,
(e.g. 10% vs 80%).
▪ S.D (𝝈) or variance: Genetic variation can be increased by hybridization, induced mutation, etc.
N.B.:(i)
Selection Value of i
intensity (%)
1 2.67
2 2.42
5 2.06
10 1.76
20 1.40
30 1.16

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(ii) When h2 is high, breeders impose higher selection pressure to achieve greater genetic advance,
but risk a rapid loss in genetic variation. For low h2, breeders exert lower selection pressure, so as
to improve on genetic gain.

(f) Calculations:
Example: Given, mean yield of F3 population from a cross of two lines of cotton = 450g/plant, Standard
deviation of the yield = 58.0g/plant, h2 of yield = 60%, the mean of the top 5% of the F3 plant
population = 570g/plant.
Calculate (i) Selection differential (S), (ii) R/∆𝐆, (iii) Yield of F4 population.
Solutions:
(i) S = 𝐗P – 𝐗I, ⟹ 570 – 450 = 120g.

𝟔𝟎
(ii) ∆𝐆 = 𝛔 . i . h2; where 𝛔 = 58.0, i at 5% = 2.06, h2 = 60% = = 0.6
𝟏𝟎𝟎
∴ ∆𝐆 = 58.0 x 2.06 x 0.6 = 71.69g.

(iii) Mean yield of F4 population = 𝐗I + ∆𝐆 = 450 + 71.69 = 521.69g/plant.

7.4 Genotype X Environment Interaction (G X E):

(a) It is the modification/influence of the genotype by the environmental factors, OR it is the failure of
genotypes to respond similarly to all environments.
● Examples,
(i) Drought may stunt all plants in the field, even if the plants had genetic constitution for tallness.
(ii) If a group of genotypes are exposed to different sets of environments, the individual genotype will
respond differently to each set of environment – as illustrated below.

E E

B B
D A
A
C C

E1 E2 E1 E2 E1 E2
No interaction No interaction

D
B
D

E1 E2 E1 E2
Interaction with magnitude Interaction with rank
changes order changes

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(b) Due to G X E, it is a common practice in plant breeding to test varieties over several locations and
years before release; to minimize error in selection, hence be sure of yield stability – in adaptation
area(s) of a cultivar. Genotypes that are less responsive to the environmental influences are more
stable in performance across varied environments, and vice versa. However, a genotype that is more
responsive to the environmental effects can be released to a smaller, specific area of adaptation.

7.0 INBREEDING DEPRESSION AND HETEROSIS

7.1 Inbreeding depression:

(a) Inbreeding: Is the mating between individuals closely related by descent or ancestry. The highest degree
of inbreeding is achieved through selfing; others – full-sib, half-sib mating.
(b) Inbreeding depression:
- It is the reduction or loss in vigour, yield and fertility of a progeny compared to parents, as a result of
inbreeding.
- The phenomenon is particularly common (detected) in cross-pollinating crop species.
- Self-pollinated species do not show significant inbreeding depression, because they have developed
homozygous balance.
* Homozygous balance: Condition where unfavourable recessive genes in homozygous state have been
eliminated, thus leaving favourable recessive homozygous genes.

7.2 Heterosis (hybrid vigour):

(a) Concept: Heterosis is the superiority of an F1 hybrid over the parents (mean value of both parents or the
highest parent value), in terms of yield or any other trait. It can be an increase or a decrease depending on
the breeding objective.
(b) Determination of:
(i) Heterosis:
𝐏𝟏 𝐏𝟐
(1) By comparing mean value of F1 with mid-parent (MP) value, i.e. ;
𝟐
𝐅𝟏 𝐌𝐏
- % heterosis = x 100
𝐌𝐏
- Average heterosis.
(2) By comparing mean value of F1 with the mean of the higher parent (HP) value;
𝐅𝟏 𝐇𝐏
- % heterosis = x 100
𝐇𝐏
- Higher or better parent heterosis.
(3) By comparing mean value of F1 with the mean of the best commercial cultivar (CC);
𝐅𝟏 𝐂𝐂
- % heterosis = x 100
𝐂𝐂
- Useful/economic heterosis.
(ii) Inbreeding depression:
𝐅𝟏 𝐅𝟐
% Inbreeding depression = x 100
𝐅𝟏
(c) In general, cross-pollinated plant species show significant heterosis, especially where genetically
divergent inbred lines are used as parents – usually exploited in commercial production of hybrid seeds.
N.B. crosses in self-pollinated species show a smaller magnitude of heterosis compared to the cross-
pollinated ones.

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(d) Manifestation of heterosis:

(i) Increased vigour, yield and fertility.
(ii) Greater adaptability, resistance to pests, etc.

7.3 Genetic basis of heterosis and inbreeding depression:

There are three theories:
(i) Dominance theory: States that heterosis is due to a large number of dominant alleles present in F 1
hybrid, e.g.
P1 X P2 → offspring
aa AA Aa
BB bb Bb
CC cc Cc
dd DD Dd
N.B. Inbreeding depression is due to the harmful effects of homozygous recessive alleles that emerge
as a result of inbreeding.
(ii) Overdominance theory: States that heterozygosity confers advantage over the corresponding
homozygosity, e.g. in aa, Aa and AA, the Aa is superior to both aa and AA, i.e. this theory equates
heterosis with heterozygosity.
(iii) Biochemical theory: Based on the physiological/biochemical process taking place in F1 and the
parents.
There are two hypotheses,
(a) Greater Initial Capital Hypothesis: Suggests that heterosis results from large embryo and
endosperm in either parent – and therefore an increase in food supply, hence increased vigour of
seedling.
(b) Cytoplasmic- Nuclear Reaction Hypothesis: Says that the interactions of the cytoplasmic and
nuclear systems from different individuals may influence hybrid vigour.