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International Journal of Advanced Engineering Technology and Innovative Science (IJAETIS)
Available online at: www.ijaetis.org
Volume1, Issue 1, Page No: 01-05
November-December 2015
ISSN:2455-1651
Corresponding author: Rajesh Wakchaure www.ijaetis.org
1
Importance of Heterosis in Animals: A Review
Rajesh Wakchaure1, Subha Ganguly2*, Praveen Kumar Praveen3, Subhash Sharma4, Avinash Kumar5, Tanvi Mahajan6 and Kausar Qadri7
1Associate Professor, Department of Animal Genetics and Breeding, 2Associate Professor and Head, Department of Veterinary Microbiology,
3Assistant Professor, Department of Veterinary Public Health and Epidemiology, 4Assistant Professor, Department of Veterinary Parasitology,
5Assistant Professor, Department of Veterinary Pharmacology and Toxicology, 6Assistant Professor, Department of Veterinary Anatomy and
Histology, 7Assistant Professor, Department of Veterinary Medicine, Arawali Veterinary College (Affiliated with Rajasthan University of Veterinary
and Animal Sciences, Bikaner), N.H. – 52 Jaipur Road, V.P.O. Bajor, Dist. Sikar, Pin – 332001, Rajasthan, India
ganguly38@gmail.com
ABSTRACT
Hybrid vigor or heterocyst is the phenomenon in which progeny of crosses between inbred lines or purebred populations
are better than the expected average of the two populations or lines for a particular trait. Utilization of heterocyst (hybrid
vigor) is the exclusive goal of crossbreeding. The amount of heterosis maintained in a herd depends on the type of
crossbreeding system selected. Hybrid vigor includes greater viability, faster growth rate, greater milk, egg and wool
production in animals. Heterocyst is an unexpected deviation from the average of the two parental lines. The cause of
heterosis the non-additive gene action (dominance, Overdominance and epistasis).No heterosis is observed for traits
governed by additive gene action. Heterosis can occur for a wide variety of performance traits. The traits showing
heterosis are called as heterotic traits. However, it tends to be greatest for traits with low heritability and least for traits
with high heritability. Traits of low heritability (reproductive traits) are generally most benefited from heterosis. They can
be improved through the adequate use of crossbreeding systems.
Keywords: crossbreeding, heterosis, crossbred
INTRODUCTION:
In 1914 Professor Shull proposed for the first time the
word ‘heterosis’ (Shull, 1914). The term used to measure
crossbred performance compared to the parental average
is hybrid vigour also known as heterosis. Hybrid Vigour
measures the ability of crossbred offspring to outperform
the expected abilities transmitted by their parents. Since
the goal of crossbreeding is to combine two, three or four
different breeds in order to achieve some desirable trait
from each different breed, Heterosis refers to the
superiority of the crossbred animal relative to the average
of its straightbred parents. Heterosis may be positive or
negative depending upon the trait. Positive heterosis is
called as hybrid vigour. Heterosis is typically expressed in
percentage improvement in the trait of interest. Heterosis
results from the increase in the heterozygosity of a
crossbred animal’s genetic makeup. Heterosis is the result
of gene dominance, overdominance and epistasis.
Heterosis dependent on an animal having two different
copies of a gene. The level of heterozygosity an animal
has depends on the random inheritance of copies of genes
from its parents. The exploitation of heterosis most
important reason for utilising cross breeding in animals
along with the exploitation of additive effects from
improved purebred animals. Heterosis arises from the
effects of gene combinations means effects of pairs of
genes (Cassell, 2007). Heterotic effects in the crossbred
progeny depends upon the differences in the frequencies
of the different alleles at each locus that contributes to the
trait (McAllister, 2002), larger these differences greater
the heterozygosity and the heterosis effects. Crossbred
animals often show increased vitality and performance.
This is known as heterosis or hybrid effect. The Sahiwal -
Friesian cross resistant to most of the common cattle
diseases and has a good milk production. A criss-cross
breeding programme is suggested to maintain the hybrid
vigour in the offspring of crossbred animals. In general,
animals that are crosses of unrelated breeds, such as
Angus and Brahman exhibit higher levels of heterosis due
to more heterozygosity than crosses of more genetically
similar breeds such as a cross of Angus and Hereford. The
genetic basis for heterosis is the opposite of the origin of
inbreeding depression. Crossbreeding cause more gene
pairs to be heterozygous. Breeds that are genetically
diverse tend to cause more heterozygosity and more
heterosis when crossed. Heterozygosity will result in
better performance if there is non-additive gene action
(dominance, overdominance and epistasis). Crossbreeding
has been shown to be an efficient method to improve
reproductive efficiency and productivity and fitness in
beef cattle. Crossbred (F1, F2 and F3) females had calves
that weighed an average of approximately 5.5 lb. more
than purebred calves at birth(Gregory et al., 1991).Cross-
breeding within species leads to offspring that are
genetically fitter than their parents (Darwin, 1876).
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Lippman and Zamir (2006) described offspring from
parents with greater genetic diversity are genetically fitter
than offspring of parents with less diversity. Cross breeds
dogs are faster learners than pure breeds (Ennik et al.,
2006). The increase in corn production (Duvick, 2001),
milk production (Ahlborn-Breier and Hokenboken,1991)
and meat production (Sellier, 1976) possible with cross-
breeding. 16% increase in the pounds of calf weaning
weight per cow exposed above the average of the parent
breeds (Ritchie et al., 1999). Crossbreeding as a mating
system optimizes the additive genetic and non-additive
(heterotic) breed effects of Bos taurus and Bos indicus
cattle in sustainable breeding systems (Gregory and
Cundiff, 1980). The reason for crossbreeding is to increase
the dairy cattle production through new combinations of
genes in different breeds (Simm, 2000). Heterosis is a
result of the non-additive gene effect, dominance and
epitasis along with differences in the frequencies of the
different alleles at each locus. The total genetic makeup of
crossbreds can include additive effects, dominance,
maternal effects, maternal heterosis and recombination
effects. Which effect that may be present is dependent of
the particular kinds of crosses involved (McAllister,
2002). The amount of heterosis expressed for a given trait
is inversely related to the heritability of the trait.
Generally, heterosis generates the largest improvement in
lowly heritable traits. Moderate improvements due to
heterosis in moderately heritable traits. Little or no
heterosis is observed in highly heritable traits. The highest
level of heterosis is most commonly seen in functional
traits affecting reproduction, survival and overall fitness.
These traits often show at least 10% heterosis and low
heritability .Production traits affecting milk yield and
growth show about 5% heterosis and a moderately high
heritability (Hansen, 2006). The expected level of
heterosis is difficult to predict and it differs depending on
the type and number of breeds in the crossbreeding system
(Sorensen et al., 2008). Crossbreeding can also cause
negative effects and one of them is recombination loss. It
is caused by separation of favorable gene combinations
that are accumulated in the parental breeds.
Recombination loss can be difficult to estimate although it
has been seen to reduce the level of heterosis (Cassell and
McAllister, 2009). The highest level of individual
heterosis is always seen in the F1 generation, but
unfortunately the level always decreases in subsequent
generations. If F1 cattle are crossed to produce the second
generation (F2), heterosis is halved compared to the level
in the F1. It continues to be halved in every following
generation of backcrossing to the parent breeds (Simm,
2000). An alternative to maintain the level of heterosis
after creating a two way cross is to produce a three way
cross because in the third generation (F3) or fourth
generation (F4) there is no further decrease in heterosis, as
long as no inbreeding exists. The level of heterosis
changes depending on the number of breeds in the cross
(Sorensen et al., 2008). Heins (2007) reported that the
Brown-Swiss-Holstein crossbreds had only a slightly
reduced milk yield along with a significantly higher yield
of milk components, fewer days open and a low number
of somatic cells compared to purebred Holstein. Due to
heterosis, crossbred Jersey-Holsteins had superior
performance compared with purebred Holsteins for milk
yield, fat and protein (Bryant et al., 2007). Jersey-Holstein
crossbred cows maintained body condition score and
hence had lower levels of live weight loss after calving
(Heins et al., 2008). Cattle and Swine species dependant
heavily on heterosis to improve productivity and
efficiency of production (Hansen, 2006). In temperate
countries, crossbreeding has been widely used in pigs and
poultry to exploit both breed differences and heterosis .
Crosses between temperate and tropical breeds have often
shown large amounts of heterosis, because of the large
genetic distance between them. Heterosis is more
important under a suboptimal (poor) than in optimal
(good) environment. Thus, heterosis is the complement of
inbreeding depression and usually appears in traits that
show depression of performance under inbreeding.
Reproductive traits in dairy cattle are usually very
sensitive to inbreeding depression and thus cross breeding
or out-crossing can show large heterotic effects. F1 crosses
probably take the advantage of hybrid vigor that arises by
crossing two genetically distant populations. Second-
generation crosses suffer reduction in hybrid vigor by half
than first generation crosses due to segregation and
recombination losses (Sendros, 2002). Loss of hybrid
vigor is the genetic factor for reduced performances in F2
crosses, poor selection standards to select F1 bulls for inter
se mating greatly contributed drop in the lifetime
performances. F1 crosses yielded more milk (147%), were
milked for more days and had shorter calving interval
(McDowell, 1988). Studies in France have shown that the
F1 crosses tend to be above median average of the two
breeds for milk but closer to the Normande for
components (Hansen, 2010). A three way cross will
maintain hybrid vigor in later generations at 86%, while a
2 way cross will at 67% and a 4 way cross at 93%
(Hansen, 2006). Three way crosses offer an increased
heterosis along with longevity, protein and fat
components, and calving ease (Snowdon, 2010). In a three
way cross program the F1 and F2 generations are both able
to maintain 100 % hybrid vigor compared to a two way
cross program where hybrid vigor drops to 50 % in the F2
generation (Pro Cross, 2009). Heterosis is utilized most
commonly in beef cattle through crossbreeding. Heterosis
in beef cattle can produce calves with enhanced
reproductive, survival, longevity (Dhuyvetter, 1998),
fertility, growth, meat quality (Peck, 2009) and disease
Rajesh Wakchaure, et al. International Journal of Advanced Engineering Technology and Innovative Science (IJAETIS)
Volume 1, Issue 2; November-December- 2015; Page No. 01-05
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resistance traits (Dandapat, 2009). The benefits of
heterosis on beef herd quality and consequently
profitability (Anderson, 1990) and herd management
programs (Brown, 2010). Heterosis achieved through
continuous crossbreeding can be used to increase weight
of calf weaned per cow exposed to breeding by 20 %
(Gregory and Cundiff, 1980). Heterosis can also increase
longevity of cows by 1.3 yr and can increase the total calf
weight weaned per cow by 30 % over the life span of a
dam (Cundiff et al., 1992). Loss of heterozygosity in inter
se mated populations does not occur if inbreeding is
evaded (Dickerson, 1973). Cows exhibit more hybrid
vigour in first and second parities than at later parities
(Cundiff et al., 1974). Herefords (Gregory and Cundiff,
1980) among the beef breeds and Holstein (McDowell,
1982) among the dairy breeds appear to have slightly
higher than average heterosis (Sorensen et al., 2008).
Heterosis has been utilized in beef production to enhance
fertility, longevity, growth and meat quality traits in
commercial herds through various cross-breeding systems.
Application of crossing systems such as three or four-
breed crosses would be very difficult, so rotational
crossbreeding systems are required to exploit breed and
heterotic effects. These schemes allow commercial
farmers to produce crossbred female replacements from
their own herds. Holstein-Friesian x Jersey crosses show
higher net income than purebred HF and Jersey cows, so
that dairy farmers mate their cows to bulls from another
breed to generate crossbred replacements with the aim of
exploiting the effects of breed and heterosis (Lopez-
Villalobos, 1998). In rotational systems heterosis is
retained at high levels, 66% in two–breed rotation, 86% in
three–breed rotation (Handley, 2001)
Measurement of Heterosis:
Percent heterosis can be calculated as:
% Heterosis = [Mean of F1 progeny- mean of parent breed] x 100
Mean of parent breed
Heterosis in F1= Mean of F1 progeny- mean of parent breed
Heterosis in F2 =1/2 heterosis in F1
Types of heterosis
There are three main types of heterosis
1) Individual heterosis: The improvement in
performance by the individual crossbred animal above
average of its parents. Examples of individual heterosis
are increased weaning weight, yearling weight and carcass
traits.
2) Maternal heterosis: Maternal heterosis is the
advantage of the crossbred mother over the average of
purebred mothers. Examples of maternal heterosis are
younger age at puberty, increased calving rate, increased
survival of her calf to weaning, pounds of calf produced in
her lifetime higher weaning weights, greater longevity in
the dam and other reproductive traits.
3) Paternal heterosis: Paternal heterosis is the advantage
of a crossbred sire over the average of purebred sires
(Buchanan, 2011). The improvement in productive and
reproductive characteristics of the bull. Examples of
paternal heterosis are reduced age at puberty,
improvements in scrotal circumference, improved sperm
concentration, increased pregnancy rate and weaning rate
when mated to cows.
Genetic basis of heterosis
There are three theories of heterosis. 1. Dominant theory
2. Over Dominance theory
3. Epistasis theory
1. Dominant theory: Superiority of hybrids to the
suppression of undesirable recessive alleles from
one parent by dominant alleles from the other
parent. The dominance hypothesis was first
expressed by the geneticist Charles Davenport
(1908)
2. Over Dominance theory: Heterozygote advantage
to the survival of many alleles those are recessive
and harmful in homozygotes. The overdominance
hypothesis was developed independently by East
(1908) and Shull (1908).
3. Epistasis theory: It postulates that gene
interactions are responsible for heterosis. The
epistasis is a phenomenon of interacting genes
which are not alleles.
Conclusion
The main benefit of crossbreeding is heterosis, which is the
improvement in genetic level in a hybrid offspring above
the average of the parent breeds. Crossbreeding schemes is
most profitable breeding strategy can assist improve
Rajesh Wakchaure, et al. International Journal of Advanced Engineering Technology and Innovative Science (IJAETIS)
Volume 1, Issue 2; November-December- 2015; Page No. 01-05
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growth, reproduction, production and maternal traits, health
and overall fitness by taking advantage of heterosis, which
results when animals from diverse backgrounds are
crossed. Inbreeding must be avoided to retain high levels of
heterozygosity and heterosis in composite breeds. The
challenge of maintaining heterosis and minimizing
inbreeding can only be met using large populations of
cattle. If no inbreeding is practiced, the heterosis is retained
in composites for several generations. F2 crosses are not
appropriate genotype of choice for dairy production
because continuous reduction in hybrid vigor. Traits of low
heritability such as fertility, milk yield and longevity are
difficult to enhance through pure breeding but are greatly
enhanced through crossbreeding leading to improvements
in survival, reproductive efficiency and growth rates. The
successful exploitation of heterosis depends upon how
superior the crosses are over the purebreds and cost of
replacement of purebred stock, therefore it is normally
practiced in poultry, swine and sheep where the fertility is
high and the cost of replacement of purebred stock is
essential.
Reference
[1] Ahlborn-Breier, G.and Hokenboken, W. D.
(1991) .Additive and non additive genetic effects
on milk production in dairy cattle: Evidence for
major individual heterosis. Journal of Daily
Science., 74: 592 – 602.
[2] Anderson, P. (1990, Jan). University of
Minnesota Extension Service. Retrieved Feb. 24,
2011, from Beef Cattle Management Update:
Crossbreeding Systems for Beef Cattle:
http://www.ansci.umn.edu/beef/beefupdates/
bcmu03.pdf.
[3] Brown, S. (2010, March 26). Crossbreds Add
Vigor to Herd. Retrieved February 7, 2011, from
Farm Journal Livestock and Production Editor:
http://www.agweb.com/article/Crossbreds_
Add_Vigor_to_Herd_193422/.
[4] Bryant, J. R., Lopez-Villalobos, N., Pryce, J. E.,
Holmes, C. W., Johnson, D. L. and Garrick, D. J.
(2007). Short communication: Effect of
environment on the expression of breed and
heterosis effects for production traits. Journal of
Dairy Science., 90(3): 1548–1553.
[5] Buchanan, D. A. (2011, Feb. 11). The Genetic
Principles of Crossbreeding. Retrieved Feb. 24,
2011., from Iowa Beef Center:
http://www.iowabeefcenter.org/Docs_cows/Gene
tic_Principles_Crossbreeding.pdf
[6] Cassell, B. (2007). Mechanisms of inbreeding
depression and heterosis for profitable dairying.
Pages 1-6 in Proc. 4th Biennial W.E. Petersen
Symposium, Univ. Minnesota, St. Paul.
[7] Cassell, B. and McAllister, J. (2009). Dairy
crossbreeding research: Results from current
projects. Virginia Cooperative Extension. 404-
494.
[8] Cundiff, L. V., Gregory, K. E. and Koch, R. M.
(1974). Effects of heterosis on reproduction in
Herford, Angus and Shorthorn cattle. J. Anim.
Sci., 38(4):711-727.
[9] Cundiff, L. V., Núñez-Dominguez, R.,
Dickerson, G. E., Gregory, K. E. and Koch, R.
M. (1992). Heterosis for lifetime production in
Hereford, Angus, Shorthorn, and crossbred cows.
J. Anim. Sci., 70:2397.
[10] Dandapat, A. (2009). Seminar on Genetic
resistance to diseases in cattle (p. 10). Pant
University of Agriculture & Technology:
http://www.authorstream.com/Presentation/anjan
_vet-190326-genetic-resistancediseases- cattle-
dandapat-sciencetechnology- ppt-powerpoint/.
[11] Darwin, C. R. (1876). The Effects of
Cross and Self-fertilization in the Vegetable
Kingdom (London: John Murray).
[12] Davenport, C. B. (1908). Degeneration,
albinism and inbreeding. Science.,28 (718):
455. doi:10.1126/science.28.718.454-
b. PMID 17771943.
[13] Dhuyvetter, J. (1998, March 5). Hybrid
Vigor PlaysKey Role in Beef Reproduction, Calf
Survival and Cow Longevity. Retrieved February
7, 2011, from NDSU Extension Service
:http://www.ext.nodak.edu/extnews/newsrelease/
1998/030598/19hybrid.htm
[14] Dickerson, G. E. (1973). Inbreeding and
heterosis in animals. In Proceedings of the
Animal Breeding and Genetics Symposium in
Honor of Dr. Jay L. Lush, pp. 54-77. American
Society of Animal Science, Champaign, IL.
[15] Gregory, K. E. and Cundiff, L. V. (1980).
Crossbreeding in beef cattle: Evaluation of
systems. Journal of Animal Science., 51:1224-
1242.
[16] Duvick, D. N. (2001). Biotechnology in
the 1930s: the development of hybrid maize.
Nature Reviews Genetics., 2: 69 – 74.
[17] East, E. M. (1908). Inbreeding in
corn. Reports of the Connecticut Agricultural
Experiments Station for 1907: 419–428.
[18] Ennik, I., Liinamo, A., Leighton, E. and
Arendonk, J. van. (2006). Suitability for field
service in 4 breeds of guide dogs. Journal of
Veterinary Behavior: Clinical Applications and
Research., 1: 67 – 74.
[19] Gregory, K. E., Cundiff, L. V. and. Koch,
R. M (1991). Breed effects and heterosis in
Rajesh Wakchaure, et al. International Journal of Advanced Engineering Technology and Innovative Science (IJAETIS)
Volume 1, Issue 2; November-December- 2015; Page No. 01-05
© 2015 IJAETIS. All Rights Reserved
Page5
Page5
Page5
Page5
Page5
Page5
Page5
Page5
Page5
Page5
Page5
Page5
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Page5
Page5
Page5
Page5
Page5
Page5
Page5
Page5
advanced generations of composite populations
for birth weight, birth date, dystocia, and survival
as traits of dam in beef cattle. J. Anim. Sci.,
69:3574-3589.
[20] Hansen, L. B. (2006). Monitoring the
worldwide genetic supply for dairy cattle with
emphasis on managing crossbreeding and
inbreeding. In: Proceedings of the 8th World
Congress of Genetics Applied to Livestock
Production, Belo Horizonte, Brasil, 01-01.
[21] Hansen, L. B. (2010)."Coopex
Montbeliarde - Montbeliarde X Holstein."
Coopex Montbéliarde – La Montbéliarde. Web.
08 Oct. 2010.
<http://www.coopex.com/croisement-
montbeliarde-xholstein-en.php>.
[22] Handley, J. (2010). Breeding Strategies
for Your Beef Herd. Retrieved Feb. 25, 2011,
from Government of Ontario: Ministry of
Agriculture Food & Rural Affairs:
http://www.omafra.gov.on.ca/english/livesto
ck/beef/facts/info_brdgstrat.htm
[23] Heins, B. J. (2007). Impact of an old
technology on profitable dairying in the 21st
Century. 4th Biennial WE Petersen Symposium,
2007.
[24] Heins, B. J., Hansen, L. B., Seykora, A.
J., Johnson, D. G., Linn, J. G., Romano, J. E. and
Hazel, A. R. (2008). Crossbreds of Jersey
Holstein compared with pure Holsteins for
production, fertility, and body and udder
measurements during first lactation. Journal of
Dairy Science., 91(3): 1270–1278.
[25] Lippman, Z. B. and Zamir, D. (2006).
Heterosis: revisiting the magic. Trends in
Genetics., 23: 60 -66
[26] López-Villalobos, N. (1998). Effects of
crossbreeding and selection on the productivity
and profitability of the New Zealand industry.
PhD thesis, Massey University, Palmerston
North, New Zealand.
[27] McAllister, A. J. (2002). Is crossbreeding
the answer to questions of dairy breed
utilization? J. Dairy Sci., 85:2352-2357.
[28] Mcdowell, R E. (1982). Crossbreeding as
a system of mating for dairy
production. Southern Co-op. Series Bull. No.
259. Louisiana Agric. Exp. Stn., Baton Rouge,
LA.
[29] McDowell, R. E. (1988). Strategies for
genetic improvement of cattle in the warm
climates. In: Second National Livestock
Improvement Conference (NLIC) held at Addis
Ababa, Ethiopia. pp. 24-26.
[30] Peck, C. (2009). Guarding Heterosis.
Retrieved Feb 7, 2011, from Yankee Creek
Ranch:
http://www.yankeecreekranch.com/images/0
1BEEFHeterosis.pdf
[31] Pro Cross. (2009). Retrieved October 9,
2010, from Creative
Genetics:http://www.creativegeneticsofca.com/pr
ocross.htm
[32] Ritchie, H., Banks, D., Buskirk, D. and
Cowley, J. (1999). Crossbreeding systems for
beef cattle. Michigan State University Extension
Bulletin, E-2701.
[33] Sellier, M. (1976). The basis of
crossbreeding in pigs: A review'. Livestock
Production Science., 3: 203- 226.
[34] Sendros, D. (2002). Genetic Factors
Affecting Milk Production, Growth, and
Reproductive Traits in Bos taurus x Bos indicus
Crosses in Ethiopia. PhD Thesis. Faculty of
Natural and Agricultural Sciences, Department of
Animal, Wildlife and Grassland Sciences.
University of Free State, Bloemfontein, South
Africa.
[35] Shull, G .H (1908). The composition of a
field of maize. Reports of the American Breeders
Association: 296–301.
[36] Shull, G. H. (1914). Duplicate genes for
capsule form in Bursa bursa-pastoris. Z. Ind.
Abstr. Ver. 12:97-149.
[37] Simm, G. (2000). Genetic Improvement
of Cattle and Sheep, 64-65, 70, 74-79, 83-95,
134-135, 201, 244-247, 354-355. Farming press.
CABI International, Wallingford, Oxon, UK.
[38] Snowdon, S. (2010).Going with a 3 way
cross an overveiw of 10 years of crossbreeding.
Www.creativegeneticsofca.com. Web. 6 Oct.
2010.
<http://www.creativegeneticsofca.com/document
s/others/australian%20crossbreeding.pdf>.
[39] Sorensen, M. K., Norberg, E., Pedersen,
J., Christensen, L. G. (2008). Invited review:
Crossbreeding in dairy cattle: A Danish
perspective. J. Dairy Sci., 91: 4116-4128