Lecture 1-1
Lecture 1-1
INSPIRING GREATNESS
Hybrid Cultivars
• Hybrid cultivars : 1st generation offspring of cross between inbred
line parents with different genotypes
– use homozygous parent lines (inbred lines) created by inbreeding (self-
fertilization)
– hybrid cultivar is different from a cultivar produced by hybridisation in self-
pollinated crops
• uniform in appearance
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Inbreeding in cross pollinated crops
• Inbreeding consists of any system of mating that leads to an increase in homozygosity
• Occurs in individuals related by ancestry when mated
• Self-fertilization leads to homozygosity – with heterozygosity being reduced by half in
each self-fertilization generation
• As homozygosity increases, genotype frequencies change but gene frequencies remain
the same
• Maize has been studied for the effects of inbreeding in cross pollinated crops
• Some crops like red clover seeds not normally obtained following self-fertilization due to
self-incompatibility
• Crops with male and female flowers on separate plants, self pollination cannot be done,
inbreeding accomplished through sib matings (mating of close relatives)
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Inbreeding
• Half-sib mating – (have one parent in common or one pollen source)
• Full-sib mating – mating between plants within the progeny of a single plant
• Sib matings increase homozygosity but homozygosity is reached more slowly than through
self-fertilization
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Consequence of inbreeding
• Major visible result of inbreeding – loss in size and vigor in progeny
• Decrease in vigor largest following the first generation of inbreeding and levels off as
homozygosity is approached (Fig 11.3).
• Decline in vigor known as inbreeding
depression
• Results from increases in the frequency of
homozygous loci with deleterious effects
• In plants with heterozygous loci, the recessive
deleterious allele is not expressed in the
plant phenotype due to masking by the
favorable dominant allele
• As homozygosity increases, many dominant
alleles are lost and the deleterious effects of
recessive alleles on the phenotype are expressed
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Effects of inbreeding: Example - forage crops
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Effects of Inbreeding – example: maize
• In cross-pollinated species such as maize – inbreeding results in:
• Reduction in plant stature
• Weak stalks and a tendency to
lodge
• increased susceptibility to
disease pathogens
• Many other deleterious plant
characteristics
• In development of inbred lines, undesirable
plants are discarded, retaining only the vigorous
plants which are self-pollinated again and the process repeated (See
figure)
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Important trait of inbred lines
• This is only expressed when the inbred line is crossed with another
inbred line and the hybrid progeny grown In a yield trial
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Hybrid Vigour or Heterosis
• Hybrid vigour – the increase in size or vigour or productivity of a hybrid
plant over the average or mean of its parents (mid-parent value)
• Another term, heterosis, was proposed by Shull to denote stimulation in size
and vigour in a hybrid as an expression of hybrid vigour
• The terms, hybrid vigour and heterosis, are synonymous and are used
interchangeably
• To be useful hybrid vigour needs to exceed the best parent in yield
productivity
• Unless a hybrid is superior to its best parent line, it has no advantage to the
breeder.
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Hybrid vigour or Heterosis
• Hybrid vigour may be manifested in many ways:
– Most often - increased vegetative growth or grain yield;
• It may also be reflected in:
– cell size, plant height, leaf size, root development, ear or head size, grain
number, seed size, biochemical pathways, etc
• Hybrid vigour – greatest when parents of the cross-pollinated crop species
are diverse
• May also be expressed among diverse genotypes of self-pollinated crops
– Detasseling
– Male sterility
– Self incompatibility.
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Hybrid Cultivars
• Basic steps in hybrid breeding
– Hybrid development.
– Hybrid evaluation
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Inbred line development
• An inbred line is a homozygous breeding line developed and maintained by
self-pollination
• The first step in making good inbred lines is choosing the correct source
material.
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Inbred line development
• Development of inbred lines (in cross-pollinated crops) usually begins with artificial self-
pollination (hand pollination) of individual S0 plants (original heterozygous selfed plant) in
a highly heterozygous and heterogeneous population.
• A progeny row of five or more S1 plants is planted from the seed of each selfed S0
plant. Plants in each row are self- pollinated by hand. Undesirable plants will be
discarded before harvest
• Methods that can be used include pedigree, bulk, single seed descent, and early
generation testing.
• The amount of inbreeding affects the vigour in inbred lines, ability to maintain their
genetic purity, the ease with which they are managed in a seed production field.
• The vigour of a line includes its ability to produce seed as a female parent or pollen as a
male parent.
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Inbred line improvement
• Backcross method often used
• Lines that excel in most attributes but lack one or few desirable
characteristics are improved by this method.
• Characteristics improved are usually highly heritable such as resistance to
diseases or insect pests.
• Quantitative characteristics that are moderately heritable such as
– maturity, plant height, seed size, and shape may also be adjusted by
backcrossing
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Evaluation for combining ability
• After the desired level of homozygosity is attained, inbred lines are evaluated
for combining ability (potential to form high-yielding hybrids) in a test cross.
• Two steps usually make up a test cross: a crossing block, then a yield trial.
• Since inbreeding causes loss of vigor (which decreases yield), yield cannot
be selected for in the inbreeding stages; breeders select for yield in a test
cross.
• Also, for characteristics of complex inheritance such as grain yield in maize,
the choice of parent material is not a clear-cut decision.
• It is helpful if inbreds to be used as parents are assessed for their capacity to
transmit certain characteristics to their progeny (combining ability).
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Evaluation for combining ability
• To evaluate many inbred lines for combining ability, a tester is used.
• There are two aspects of combining ability :
– general combining ability (GCA) which applies to all crosses, and
– specific combining ability (SCA) which refers to certain specific crosses
only.
• The type of tester to use for the evaluation of inbred lines in hybrid
combination depends primarily on whether information is desired on
GCA or SCA.
INSPIRING GREATNESS
General Combining Ability
• General combining ability (GCA) is the average performance of an inbred line in a number of hybrid
combinations in comparison to the contribution of other inbred lines in the same series of hybrid
combinations
• Provides information on which inbred lines should produce the best hybrids when crossed with many other
lines.
• Evaluated by crossing a line with other inbred lines and comparing the overall performance of the single-
cross progenies.
• It evaluates the additive portion of the genetic effects
• Example:
– inbred lines A, B, C , D, E crossed in all possible combinations (diallel mating)
– Single cross hybrids evaluated in yield trials
– The inbred whose single cross has the highest average yield would have the greatest GCA
• Number of possible single-cross hybrids in a half-diallel = n(n-1)/2: where n = number of inbreds
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General Combining Ability
• With large numbers of inbred lines – not feasible to conduct diallel mating
• Mixture of pollen used to pollinate the inbred lines (e.g from open-pollinated variety, double-
cross or single-cross hybrid) – highly correlated with the average performance of the inbred
line in different single-cross combinations
• Testers should be selected for their ability to determine which lines will combine well with
many other lines.
• Pollinating new inbred lines with heterogenous genotype proved to be an efficient tester
• Testcross progeny evaluated in yield trials and only inbred lines with superior GCA retained
for testing single cross combinations
INSPIRING GREATNESS
Specific Combining Ability
• Specific combining ability (SCA) is the contribution of an inbred line to hybrid
performance in a cross with a specified inbred line, in relation to its
contributions in crosses with an array of specified inbred lines.
• In other words, it is the individual performance of an inbred line in a single
specific hybrid combination.
• It evaluates non-additive gene action (dominance, epistasis, digenic
interactions)
• It is utilized to identify inbred x inbred cross combination with superior
performance
• Cross inbred lines with superior GCA in all possible combinations (diallel
crossing)
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Specific Combining Ability
• Example: A, B, C, D, E inbred lines, combination A x E produces highest single
cross yield performance, thus A x E has superior SCA
• Single-cross performance depends on favourable genes from each of the inbred
lines used in the cross
• Inbreds from unrelated populations – produce high-yielding single crosses more
frequently than inbreds from related material
• Information on specific combining ability may not provide reliable information
on the relative usefulness of an inbred line when crossed with other testers.
• Desirable single cross seed parents make excellent testers for determining
specific combining ability for three way crosses.
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Example: Test-Cross (Line x Tester) Mating Design
• Choice of testers is important
• Both natural and artificial crossing can be used for test crossing in many
crop species
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Test-Cross (Line x Tester) Mating Design
• In the case of natural hybridization, an isolated crossing block is required.
• The tester is used as the male parent and the line or clone as the female
parent.
• The lines to be tested should therefore either be
– male-sterile or self-incompatible
• The ratio of the number of male rows to female rows varies with crop species.
• Artificial crossing is used if isolation is not possible.
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Broad- versus narrow-based testers
• Testers should be selected for their ability to determine which lines will combine well
with many other lines.
• Because of their heterogeneity, open-pollinated varieties and synthetics are usually used
for determining general combining ability.
• Narrow-based testers - elite inbred lines, related inbreds lines, single crosses of related
lines
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Diallel mating design
• The diallel cross can be defined as all crosses among a
group of parents including the parents themselves
• Produce the most accurate results and provide the most
genetic information
• Differ according to whether the parents selfed and/or the
reciprocal F1’s included.
• If reciprocals are excluded – diallel is termed half diallel
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Example 2: Diallel Mating Design
• Full diallel cross or half-diallel
Females/males 1 2 3 4 5 means
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Methods of diallel crossing proposed by Griffing, 1956
Method Characteristics Number of crosses
1. or Full diallel Parents (n*), F1 and reciprocals are n2 crosses
included
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Three—way hybrids
• Produced by combining a single-cross
o and an inbred line.
• Two inbred lines crossed initially to give an F1. The F1 is then crossed with another inbred line.
• The single cross (F1) is highly productive and serves as female and the inbred line serves as male
• Three-way cross hybrids yield less than single cross hybrids.
• An added step in the seed production process
• Less uniform than single-cross hybrids
• Maintained as inbred lines
INSPIRING GREATNESS
Double cross hybrids
• Double- cross hybrids are produced by crossing two different
single crosses, giving the pedigree [(AxB)x(CxD).].
• The female is a highly productive single- cross and the male is
also a single-cross that produces pollen abundantly.
• Double cross hybrid seed production is a practical and
economical way of producing adequate seeds for farmers
• Less cost of seed production because both parents are
hybrids
• Double cross hybrids yield less than three-way hybrids
• Maintained as inbred lines
• More variable than single or 3-way crosses thus allowing
breeders to bring more different desirable characteristics
together into one hybrid than is possible in a single cross.
INSPIRING GREATNESS
Top Cross hybrids
• One of the parents is an open-pollinated variety and the other is a
single-cross hybrid or an inbred line.
• Top- cross hybrids are very useful at the initial stages of hybrid seed
production when seed companies are still learning the rudiments of
hybrid seed production.
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Hybrid characteristics
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Summary of factors that should be considered
during the production and marketing of various
types of hybrids
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Open-Pollinated Varieties
• Open-pollination – occurs in cross-pollinated species where pollination occurs naturally
and randomly without restriction.
Important attributes of open-pollinated cultivars
• Easier seed multiplication and less costly.
• Broad-based (2 to many parents used)
• Have considerable genetic buffering capacity to withstand biotic and abiotic stresses.
• Stable yields; an OPV consists of plants of different genetic makeup (genotypes) that
respond differently to environmental stresses (e.g., moisture stress, diseases,
temperature, etc.) and, as a result, it is more tolerant to stress than a hybrid.
• Thus, OPVs are particularly useful in stress prone environments.
• Cultivars show more variation for important agronomic characteristics.
INSPIRING GREATNESS
Open-Pollinated Varieties
• Seed is less expensive
• Seed can be retained and planted from season to season without suffering
considerable yield depression.
• Seed replacement is after 3 -4 years.
• Exhibit less heterosis – lower yields.
• Varietal maintenance is easier.
• F1 to F4 seed is planted.
• Communities can produce certified seed of OPVs.
• Essential to buy certified seed of an improved OPV every three to five years
in order to maintain its genetic purity.
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Hybrids vs Open-Pollinated varieties
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Heterotic relationships/patterns
• The choice of germplasm is critical in hybrid breeding
• Selection of parents may involve consideration of heterotic relationships among available
genotypes
• Performance of a hybrid is a function of the amount of heterosis expressed in the cross
between two parents.
• Heterosis generally increases as the genetic diversity between the parents increases.
• Therefore, breeders of hybrid cultivars sometimes attempt to maintain two germplasm
pools (heterotic pools) that provide desired heterosis in crosses between them.
• Thus, heterotic pools should be highly heterotic. Heterotic pools should also be broad
based.
• It is important that heterotic pools are made as divergent as possible.
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Heterotic relationships/patterns
• The probability of identifying superior crosses is greater between populations (germplasm
pools) of known heterotic patterns than within a population.
• Heterotic patterns have a strong impact on a breeding programme and once established
are not easy to change
• When you get new germplasm to enhance variation in your breeding programme, these
materials should be testcrossed to two or more heterotic stocks (testers).
• The entries showing heterosis with a tester are assigned to the opposite heterotic pool.
• It is more desirable to work with inbred lines when classifying germplasm into heterotic
pools.
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Heterosis
• Heterosis defined as the superiority of an F1 hybrid
over both of its parents in terms of yield or some
other character. o
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Heterosis
• In some cases, heterosis estimated in relation to the best commercial variety grown
since commercial usefulness of a hybrid would primarily depend on its performance in
comparison to the best commercial variety of the concerned species.
• There are few cases where F1 hybrids are inferior to their parents, i.e., hybrid vigour in
the negative direction.
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Manifestation of heterosis
• Increased yield
• Greater adaptability
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Measurement of Heterosis
Mid-parent heterosis (%) = F1 − MP ×100
MP
This refers to heterosis over the standard check variety to show the superiority of
the hybrid over the prevalent variety.
F1 = performance of hybrid
Where,
MP = average performance of parents per se
HP = performance of best parent
CV = performance of standard check variety.
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Reproduction Systems, Propagation
Methods and Types of Varieties
Type of variety/ (Modified from
Natural reproduction Breeding Category Becker, 1993)
Propagation
system process
Self-fertilization Self-fertilization Line varieties
Controlled crossing
Partially autogamous Hybrid, synthetic
Sexual
of parental
Partially allogamous varieties
components
Open pollinated
Cross-fertilization Open pollination
varieties
Vegetative
Clonal varieties
Asexual
propagation
• Differ in the degree of genetic variability maintained within the variety, in the degree of heterozygosity and in the easy with which they
can be reproduced. INSPIRING GREATNESS
Genetic Variation within a variety
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Genetic structure of varieties
• Pure line varieties of self-pollinated crops are homogeneous and homozygous and could
theoretically just be made up of one single genotype. That can easily be reproduced.
• Single-cross hybrids may also be made up of only one genotype (the offspring from a cross between
two homozygous parental lines). However, they have very high degree of heterozygosity and cannot be
easily reproduced by farmers.
• Other types of hybrids will have different levels of diversity within them, e.g., top cross hybrids where one
parent is an open-pollinated variety of a cross-pollinated crop.
• Open-pollinated varieties have a high degree of intra-varietal diversity. Heterozygosity is also present in
such varieties, depending on the outcrossing rate of the crop and the diversity of alleles for genes in the
population. Open-pollinated varieties can be reproduced easily if contamination with pollen from other
varieties can be prevented.
• Variety mixtures (multi-line varieties) of some landraces of self-pollinating crops maybe both
homozygous and heterogeneous. They are reproducible if natural selection pressures do not differ very
much from the conditions under which they are developed, so that specific types or components will not
disappear.
INSPIRING GREATNESS