Heterosis

Heterosis is ubiquitous and is of great practical value in plant breeding. The commercial
exploitation of heterosis or hybrid vigor through the development and cultivation of hybrid
cultivars is one of the landmark achievements in plant breeding. Ever since the two pioneering
publications by George H. Shull more than century years ago, in which he scientifically described
heterosis and laid the foundation of modern hybrid breeding in maize, the exploitation of heterosis
in crop and tree species has greatly expanded and the acreage under hybrid cultivars has steadily
increased. Thus, hybrid breeding has made commendable contributions in meeting the food, feed,
and fiber needs of the increasing population of the world, and benefitted farmers and consumers.

How Is Heterosis Defined?

Heterosis or hybrid vigor is the tendency of outbred strains to exceed both inbred parents in fitness.

It can also be defined as “Heterosis is the situation in which hybrid offspring exhibit phenotypic
performance that is superior to that of their
parents.”

The conceptual opposite of heterosis is inbreeding

depression. This is the loss of vigor following
related mattings. Heterosis is often viewed as
maximizing heterozygosity and, in contrast,
inbreeding depression is due to reduction in
heterozygosity.

The amount of heterosis in any hybrid relative to its

parents is trait-dependent and hybrids cannot be
simply classified as heterotic or nonheterotic. From
a research standpoint, this indicates that the search
for mechanisms of heterosis must be conducted
within the biological context of specific traits; in a practical context, it motivates research to better
predict heterotic hybrids that will provide maximum productivity for specific traits of interest.
Genetic Basis of the Heterosis:
The difficulty in formulating the genetic basis of heterosis has at least two major contributors.
First, in most cases, multiple genes contribute to the response of the F1 hybrid. Thus, sorting
through the contributions of the responsible factors is not an easy task. Secondly, the multiple
genes interact in ways that mask the action of each other in the process of epistasis. Thus, not only
do multiple genes present a complicating factor, but also the shifting states of heterozygosity or
homozygosity of individual factors can influence the impact of other genes.

Two competing hypotheses were developed to explain the heterosis;

Dominance hypothesis:

The dominance hypothesis attributes the superiority of hybrids to the suppression of undesirable
recessive alleles from one parent by dominant alleles from the other. It attributes the poor
performance of inbred strains to loss of genetic diversity, with the strains becoming purely
homozygous at many loci. The dominance hypothesis was first expressed in 1908 by the
geneticist Charles Davenport. Under the dominance hypothesis, deleterious alleles are expected to
be maintained in a random-mating population at a selection–mutation balance that would depend
on the rate of mutation, the effect of the alleles and the degree to which alleles are expressed in
heterozygotes.

Overdominance hypothesis:

Certain combinations of alleles that can be obtained by crossing two inbred strains
are advantageous in the heterozygote. The overdominance hypothesis attributes the heterozygote
advantage to the survival of many alleles that are recessive and harmful in homozygotes. It
attributes the poor performance of inbred strains to a high percentage of these harmful recessives.
The overdominance hypothesis was developed independently by Edward M.
East (1908)[6] and George Shull (1908). Genetic variation at an over dominant locus is expected to
be maintained by balancing selection.
More recently, an epigenetic component of hybrid vigor has also been established.

Epigenetic hypothesis:

The role of epistasis in heterotic and nonheterotic trait performance remains intriguing and
perplexing. Conceptually, many and diverse complex pathways interact to produce phenotypes in
individuals supporting the likelihood that genetic epistasis should be detected. However, genetic
epistasis requires not only interacting molecular pathways, but also allelic variation within
interacting pathways of enough magnitude to provide a significant statistical interaction. Large
QTL mapping studies find little evidence for epistatic interactions for specific developmental,
architectural, and biochemical traits although, as described previously, heterosis is greater for
highly complex traits such as grain yield, traits for which quantitative genetic studies more often
support the role of epistasis.
Methods for Estimation of Heterosis:
Heterosis is estimated in three different ways,

 Mid parent heterosis

 Better parent heterosis
 Commercial heterosis

Mid Parent Heterosis:

When the heterosis is estimated over the mid parent i.e. mean value or average of the two parents
is known as mid parent heterosis. It is also known as average heterosis or relative heterosis and
calculated by using formula.

F1− MP
Mid Parent Heterosis = x 100
𝑀𝑃

Where F1 is mean of F1 and MP is mean of two parents.

Better Parent Heterosis:

When the heterosis is estimated over the better parent is known as better parent heterosis. It is also
known as heterobeltiosis and calculated by using formula:
 F1− BP
Better Parent Heterosis = x 100
𝐵𝑃

The term heterobeltiosis was used by Bitzer et al (1968) to describe the improvement of
heterozygote over the better parent of the cross.

Commercial heterosis:
It refers to the superiority of F1 over the standard commercial check variety. It is also called as
economic heterosis or useful heterosis and calculated by using formula.

F1− CV
Commercial Heterosis = x 100
𝐶𝑉

Heterosis leads to increase in yield, reproductive ability, adaptability, disease and insect resistance,
general vigour, quality etc. For most of the characters, the desirable heterosis is positive. But for
some characters like earliness, height in cereals and toxic substances are negative heterosis.
 Combining Ability
Identification of the best performing lines (for commercial release) and lines which can be used as
parents in future crosses are two principal objects considering in most crop breeding programs.
The best performing lines for required characteristics are selected based on conducting multi-
environment trials following statistical analysis.

What is Combining Ability:

Crossing a line to several others provides the mean performance of the line in all its crosses.
Combining ability or productivity in crosses is defined as the cultivars or parents’ ability to
combine among each other during hybridization process such that desirable genes or characters
are transmitted to their progenies.

At first, combining ability was a general concept used collectively for classifying an inbred line
respective to its cross performance but was later amended. Sprague and Tatum defined GCA as
the average performance of a genotype in a series of hybrid combinations. They defined SCA
as those cases in which certain hybrid combinations perform better or poorer than would be
expected based on the average performance of the parental inbred lines. Parents showing a
high average combining ability in crosses are considered to have good GCA while if their potential
to combine well is bounded to a cross, they are considered to have good SCA.

From a statistical point of view, the GCA is a main effect and the SCA is an interaction effect.
Based on Sprague and Tatum, GCA is owing to the activity of genes which are largely additive in
their effects as well as additive × additive interactions. Specific combining ability is regarded as
an indication of loci with dominance variance (non-additive effects) and all the three types of
epistatic interaction components if epistasis were present. They include additive × dominance and
dominance × dominance interactions.

Importance of GCA and SCA:

GCA is an effective tool used in selection of parents based on performance of their progenies,
usually the F1 but it has also been used in F2 and later generations .A high GCA value shows that
the parental mean is superior or inferior to the general mean. This indicates a potent evidence of
desirable gene flow from parents to offspring at high intensity and represents information
regarding the concentration of predominantly additive genes. A high GCA estimate indicates
higher heritability and less environmental effects. It may also result in less gene interactions and
higher achievement in selection.

Observations of performance of different cross patterns based on SCA have been used to make
inferences on gene action at play. High SCA effects resulting from crosses where both parents are
good general combiners (i.e., good GCA × good GCA) may be ascribed to additive × additive gene
action. The high SCA effects derived from crosses including good × poor general combiner parents
may be attributed to favorable additive effects of the good general combiner parent and epistatic
effects of poor general combiner, which fulfils the favorable plant attribute. High SCA effects
manifested by low × low crosses may be due to dominance × dominance type of non-allelic gene
interaction producing over dominance thus being non-fixable.

Different methods have been used to evaluate relative importance of GCA and SCA in plant
breeding. The first step is to check whether both GCA and SCA are significant at P=0.05 or at
higher probability levels (0.01 or 0.001 etc.). If both the GCA and SCA values are not significant,
epistatic gene effects may play a remarkable role in determining these characters.

The ratio of combining ability variance components (predictability ratio) determines the type of
gene action involved in the expression of traits and allows inferences about optimum allocation of
resources in hybrid breeding:

σ2 GCA
σ2 SCA
in which σ2gca refers to general combining ability variance and σ2sca refers to specific combining
ability variance. The closer this ratio is to one, the greater the prediction of GCA alone, whereas a
ratio with a value less than 1 shows SCA action.
Estimation of combining Ability:

References:
Fasahat and Parviz. 2016. Principles and Utilization of Combining Ability in Plant Breeding.
Biometrics & Biostatistics International Journal. 4.

Hochholdinger. 2007. Towards the molecular basis of heterosis. Trends in plant science. 12. 427-
32.

Kaeppler, S. 2011. Heterosis: One boat at a time, or a rising tide?. The New Phytologist. 189(4).
900-902.

Schnable and Nathan. 2013. Progress Toward Understanding Heterosis in Crop Plants. Annual
review of plant biology. 64.

Li, C., Zhao and T., Yu 2018. Genetic basis of heterosis for yield and yield components explored
by QTL mapping across four genetic populations in upland cotton. BMC
Genomics 19. 910.