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Heterosis, hybrid vigor, or outbreeding enhancement, is the improved or increased function of any biological quality in a hybrid offspring. The adjective derived from heterosis is heterotic.

An offspring exhibits heterosis if its traits are enhanced as a result of mixing the genetic contributions of its parents. These effects can be due to Mendelian or non-Mendelian inheritance.

Definitions

In proposing the term heterosis to replace the older term heterozygosis, G.H. Shull aimed to avoid limiting the term to the effects that can be explained by heterozygosity in Mendelian inheritance.^[1]

The physiological vigor of an organism as manifested in its rapidity of growth, its height and general robustness, is positively correlated with the degree of dissimilarity in the gametes by whose union the organism was formed … The more numerous the differences between the uniting gametes — at least within certain limits — the greater on the whole is the amount of stimulation … These differences need not be Mendelian in their inheritance … To avoid the implication that all the genotypic differences which stimulate cell-division, growth and other physiological activities of an organism are Mendelian in their inheritance and also to gain brevity of expression I suggest … that the word 'heterosis' be adopted.

Heterosis is often discussed as the opposite of inbreeding depression although differences in these two concepts can be seen in evolutionary considerations such as the role of genetic variation or the effects of genetic drift in small populations on these concepts. Inbreeding depression occurs when related parents have children with traits that negatively influence their fitness largely due to homozygosity. In such instances, outcrossing should result in heterosis.

Not all outcrosses result in heterosis. For example, when a hybrid inherits traits from its parents that are not fully compatible, fitness can be reduced. This is a form of outbreeding depression.

Controversy

The term heterosis often causes confusion and even controversy, particularly in selective breeding of domestic animals, because it is sometimes claimed that all crossbred plants and animals are "genetically superior" to their parents, due to heterosis^{[citation needed]}. However, there are two problems with this claim:

First, "genetic superiority" is an ill-defined term and not generally accepted terminology within the scientific field of genetics.^[2] A related term fitness is well defined, but it can rarely be directly measured. Instead, scientists use objective, measurable quantities, such as the number of seeds a plant produces, the germination rate of a seed, or the percentage of organisms that survive to reproductive age.^[3] Within this perspective, crossbred plants and animals exhibiting heterosis may have "superior" production on these scales, but this does not necessarily equate to evidence of "genetic superiority". Use of the term "genetic superiority" is a value judgement, generally in the realm of politics, and is not science.^[2]
Second, not all hybrids exhibit heterosis (see outbreeding depression).

A clearly ambiguous counter-example to any value judgement on hybrids and hybrid vigor is the mule. While mules are almost always infertile, they are valued for a combination of hardiness and temperament that is different from either of their horse or donkey parents. While these qualities may make them "superior" for particular uses by humans, the infertility issue implies that these animals would most likely become extinct without the intervention of humans through animal husbandry, making them "inferior" in terms of natural selection.

Some modern geneticists refrain from even using the terms inferior and superior due to the association of these words with political movements that espouse genocide.

Genetic and epigenetic bases of heterosis

Since the early 1900s (as discussed in the article Dominance versus overdominance) two competing genetic hypotheses, not necessarily mutually exclusive, have been developed to explain hybrid vigor. More recently, an epigenetic component of hybrid vigor has also been established.^[4]^[5]

The genetic dominance hypothesis attributes the superiority of hybrids to the masking of expression of undesirable (deleterious) recessive alleles from one parent by dominant (usually wild-type) alleles from the other (see Complementation (genetics)). It attributes the poor performance of inbred strains to the expression of homozygous deleterious recessive alleles. The genetic overdominance hypothesis states that some combinations of alleles (which can be obtained by crossing two inbred strains) are especially advantageous when paired in a heterozygous individual. This hypothesis is commonly invoked to explain the persistence of some alleles (most famously the Sickle cell trait allele) that are harmful in homozygotes. In normal circumstances, such harmful alleles would be removed from a population through the process of natural selection. Like the dominance hypothesis, it attributes the poor performance of inbred strains to expression of such harmful recessive alleles. In any case, outcross matings provide the benefit of masking deleterious recessive alleles in progeny. This benefit has been proposed to be a major factor in the maintenance of sexual reproduction among eukaryotes, as summarized in the article Evolution of sexual reproduction.

An epigenetic contribution to heterosis has been established in plants,^[5] and it has also been reported in animals.^[6] MicroRNAs (miRNAs), discovered in 1993, are a class of non-coding small RNAs which repress the translation of messenger RNAs (mRNAs) or cause degradation of mRNAs.^[7] In hybrid plants, most miRNAs have non-additive expression (it might be higher or lower than the levels in the parents).^[5] This suggests that the small RNAs are involved in the growth, vigor and adaptation of hybrids.^[5]

It was also shown^[4] that hybrid vigor in an allopolyploid hybrid of two Arabidopsis species was due to epigenetic control in the upstream regions of two genes, which caused major downstream alteration in chlorophyll and starch accumulation. The mechanism involves acetylation and/or methylation of specific amino acids in histone H3, a protein closely associated with DNA, which can either activate or repress associated genes.

MHC in animals

One example of where particular genes may be important in vertebrate animals for heterosis is the major histocompatibility complex. Vertebrates inherit several copies of both MHC class I and MHC class II from each parent, which are used in antigen presentation as part of the adaptive immune system. Each different copy of the genes is able to bind and present a different set of potential peptides to T-lymphocytes. These genes are highly polymorphic throughout populations, but will be more similar in smaller, more closely related populations. Breeding between more genetically distant individuals will decrease the chance of inheriting two alleles which are the same or similar, allowing a more diverse range of peptides to be presented. This therefore gives a decreased chance that any particular pathogen will not be recognised, and means that more antigenic proteins on any pathogen are likely to be recognised, giving a greater range of T-cell activation and therefore a greater response. This will also mean that the immunity acquired to the pathogen will be against a greater range of antigens, meaning that the pathogen must mutate more before immunity is lost. Thus hybrids will be less likely to be succumb to pathogenic disease and will be more capable of fighting off infection.

In plants

Crosses between inbreds from different heterotic groups result in vigorous F1 hybrids with significantly more heterosis than F1 hybrids from inbreds within the same heterotic group or pattern. Heterotic groups are created by plant breeders to classify inbred lines, and can be progressively improved by reciprocal recurrent selection.

Heterosis is used to increase yields, uniformity, and vigor. Hybrid breeding methods are used in maize, sorghum, rice, sugar beet, onion, spinach, sunflowers, broccoli and to create a more psychoactive cannabis.

Corn (maize)

Nearly all field corn (maize) grown in most developed nations exhibits heterosis. Modern corn hybrids substantially outyield conventional cultivars and respond better to fertilizer.

Corn heterosis was famously demonstrated in the early 20th century by George H. Shull and Edward M. East after hybrid corn was invented by Dr. William James Beal of Michigan State University based on work begun in 1879 at the urging of Charles Darwin. Dr. Beal's work led to the first published account of a field experiment demonstrating hybrid vigor in corn, by Eugene Davenport and Perry Holden, 1881. These various pioneers of botany and related fields showed that crosses of inbred lines made from a Southern dent and a Northern flint, respectively, showed substantial heterosis and outyielded conventional cultivars of that era. However, at that time such hybrids could not be economically made on a large scale for use by farmers. Donald F. Jones at the Connecticut Agricultural Experiment Station, New Haven invented the first practical method of producing a high-yielding hybrid maize in 1914-1917. Jones' method produced a double-cross hybrid, which requires two crossing steps working from four distinct original inbred lines. Later work by corn breeders produced inbred lines with sufficient vigor for practical production of a commercial hybrid in a single step, the single-cross hybrids. Single-cross hybrids are made from just two original parent inbreds. They are generally more vigorous and also more uniform than the earlier double-cross hybrids. The process of creating these hybrids often involves detasseling.

Temperate maize hybrids are derived from two main heterotic groups: Iowa Stiff Stalk Synthetic, often referred to as BSSS,^{[clarification needed]} and non stiff stalk.^{[citation needed]}

Rice (Oryza sativa)

Rice production has seen enormous rise in China due to heavy uses of hybrid rice. In China, efforts have generated a super hybrid rice strain (LYP9) with a production capability of ~15 tons per hectare. In India also, several varieties have shown high vigor, including RH-10 and Suruchi 5401.

Hybrid livestock

The concept of heterosis is also applied in the production of commercial livestock. In cattle, hybrids between Black Angus and Hereford produce a hybrid known as a "Black Baldy". In swine, "blue butts" are produced by the cross of Hampshire and Yorkshire. Other, more exotic hybrids such as "beefalo" are also used for specialty markets.

Within poultry, sex-linked genes have been used to create hybrids in which males and females can be sorted at one day old by color. Specific genes used for this are genes for barring and wing feather growth. Crosses of this sort create what are sold as Black Sex-links, Red Sex-links, and various other crosses that are known by trade names.

Commercial broilers are produced by crossing different strains of White Rocks and White Cornish, the Cornish providing a large frame and the Rocks providing the fast rate of gain. The hybrid vigor produced allows the production of uniform birds with a marketable carcass at 6–9 weeks of age.

Likewise, hybrids between different strains of White Leghorn are used to produce laying flocks that provide the majority of white eggs for sale in the United States.

Humans

Any evidence of heterosis in humans is derived from observational studies.^[8] It has been suggested that increased average height as well as many beneficial effects on average health and intelligence have resulted from an increased heterosis, resulting from increased mixing of the human population such as by urbanization.^[9] Diverse human populations commonly have a cultural rule or norm that prohibits sexual relations between relatives. Such a prohibition is referred to as an incest taboo. An effect of the incest taboo is the promotion of heterosis and avoidance of congenital birth defects that often result from expression of deleterious recessive alleles in children from matings between close relatives.

References

^ George Harrison Shull (1948). "What Is "Heterosis"?". Genetics. 33 (5): 439–446. PMC 1209417. PMID 17247290.
^ ^a ^b Risch N, Burchard E, Ziv E, Tang H (July 2002). "Categorization of humans in biomedical research: genes, race and disease". Genome Biol. 3 (7): comment2007. doi:10.1186/gb-2002-3-7-comment2007. PMC 139378. PMID 12184798.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
^ Weller SG, Sakai AK, Thai DA, Tom J, Rankin AE (November 2005). "Inbreeding depression and heterosis in populations of Schiedea viscosa, a highly selfing species". J. Evol. Biol. 18 (6): 1434–44. doi:10.1111/j.1420-9101.2005.00965.x. PMID 16313456.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b Ni Z, Kim ED, Ha M; et al. (January 2009). "Altered circadian rhythms regulate growth vigour in hybrids and allopolyploids". Nature. 457 (7227): 327–31. doi:10.1038/nature07523. PMC 2679702. PMID 19029881. {{cite journal}}: Explicit use of et al. in: |author= (help)CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c ^d Baranwal VK, Mikkilineni V, Zehr UB, Tyagi AK, Kapoor S (November 2012). "Heterosis: emerging ideas about hybrid vigour". J. Exp. Bot. 63 (18): 6309–14. doi:10.1093/jxb/ers291. PMID 23095992.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Han Z, Mtango NR, Patel BG, Sapienza C, Latham KE (October 2008). "Hybrid vigor and transgenerational epigenetic effects on early mouse embryo phenotype". Biol. Reprod. 79 (4): 638–48. doi:10.1095/biolreprod.108.069096. PMC 2844494. PMID 18562704.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Zhou Y, Ferguson J, Chang JT, Kluger Y (2007). "Inter- and intra-combinatorial regulation by transcription factors and microRNAs". BMC Genomics. 8: 396. doi:10.1186/1471-2164-8-396. PMC 2206040. PMID 17971223.{{cite journal}}: CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)
^ Attention: This template ({{cite pmid}}) is deprecated. To cite the publication identified by PMID 13266803, please use {{cite journal}} with |pmid=13266803 instead.
^ Last paragraphs in Discussion section in: Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi:10.1038/ejhg.2008.48, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi=10.1038/ejhg.2008.48 instead.

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 == Humans ==
 {{See also|Miscegenation}}
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 Any evidence of heterosis in humans is derived from [[observational study|observational studies]].<ref>{{cite pmid|13266803}}</ref> It has been suggested that increased average height as well as many beneficial effects on average health and intelligence have resulted from an increased heterosis, resulting from increased mixing of the [[human population]] such as by [[urbanization]].<ref>Last paragraphs in ''Discussion'' section in: {{cite doi|10.1038/ejhg.2008.48}}</ref> Diverse human populations commonly have a cultural rule or norm that prohibits sexual relations between relatives.  Such a prohibition is referred to as an [[incest taboo]].  An effect of the incest taboo is the promotion of heterosis and avoidance of congenital birth defects that often result from expression of deleterious recessive alleles in children from matings between close relatives.