Hybrid (biology) - Can Hybrid Animals Reproduce

In biology a hybrid, also known as cross breed, is the result of mixing, through sexual reproduction, two animals or plants of different breeds, varieties, species or genera. Using genetic terminology, it may be defined as follows:

1. Hybrid generally refers to any offspring resulting from the breeding of two genetically distinct individuals, which usually will result in a high degree of heterozygosity, though hybrid and heterozygous are not, strictly speaking, synonymous;
2. genetic hybrid carries two different alleles of the same gene;
3. structural hybrid results from the fusion of gametes that have differing structure in at least one chromosome, as a result of structural abnormalities;
4. numerical hybrid results from the fusion of gametes having different haploid numbers of chromosomes;
5. permanent hybrid is a situation where only the heterozygous genotype occurs, because all homozygous combinations are lethal.

From a taxonomic perspective, hybrid refers to:

1. Offspring resulting from the interbreeding between two animal species or plant species. See also hybrid speciation.
2. Hybrids between different subspecies within a species (such as between the Bengal tiger and Siberian tiger) are known as intra-specific hybrids. Hybrids between different species within the same genus (such as between lions and tigers) are sometimes known as interspecific hybrids or crosses. Hybrids between different genera (such as between sheep and goats) are known as intergeneric hybrids. Extremely rare interfamilial hybrids have been known to occur (such as the guineafowl hybrids). No interordinal (between different orders) animal hybrids are known.
3. The third type of hybrid consists of crosses between populations, breeds or cultivars within a single species. This meaning is often used in plant and animal breeding, where hybrids are commonly produced and selected, because they have desirable characteristics not found or inconsistently present in the parent individuals or populations.

Terminology

The term hybrid is derived from Latin hybrida, meaning the "offspring of a tame sow and a wild boar", "child of a freeman and slave", etc. The term came into popular use in English in the 19th century, though examples of its use have been found from the early 17th century.

There is a popular convention of naming hybrids by forming portmanteau words. This became common in the 1920s, with the breeding of tiger-lion hybrids (liger and tigon). This was playfully (but unsystematically) extended to a number of other hybrids, or hypothetical hybrids, such as beefalo (1960s), humanzee (1980s), and cama (1998).

Types of hybrids

Depending on the parents, there are a number of different types of hybrids;

• Single cross hybrids  result from the cross between two true breeding organisms which produces an F1 generation called an F1 hybrid (F1 is short for Filial 1, meaning "first offspring"). The cross between two different homozygous lines produces an F1 hybrid that is heterozygous; having two alleles, one contributed by each parent and typically one is dominant and the other recessive. Typically, the F1 generation is also phenotypically homogeneous, producing offspring that are all similar to each other.
• Double cross hybrids result from the cross between two different F1 hybrids.
• Three-way cross hybrids result from the cross between an F1 hybrid and an inbred line.
• Triple cross hybrids result from the crossing of two different three-way cross hybrids.
• Population hybrids result from the crossing of plants or animals in a population with those of another population. These include crosses between organisms such as interspecific hybrids or crosses between different breeds.
• Stable hybrid is a horticultural term, usually referring to an annual plant that, if grown and bred in a small monoculture free of external pollen (e.g., an air-filtered greenhouse) will produce offspring that are "true to type" with respect to phenotype; i.e., a true breeding organism.
• Hybrid species result from hybrid populations evolving reproductive barriers against their parent species through hybrid speciation.

Interspecific hybrids

Interspecific hybrids are bred by mating two species, normally from within the same genus. The offspring display traits and characteristics of both parents. The offspring of an interspecific cross are often sterile; thus, hybrid sterility prevents the movement of genes from one species to the other, keeping both species distinct. Sterility is often attributed to the different number of chromosomes between the two species. For example, donkeys have 62 chromosomes, while horses have 64 chromosomes, and mules and hinnies have 63 chromosomes. Mules, hinnies, and other normally sterile interspecific hybrids cannot produce viable gametes, because differences in chromosome structure prevent appropriate pairing and segregation during meiosis, meiosis is disrupted, and viable sperm and eggs are not formed. However, fertility in female mules has been reported with a donkey as the father.

Most often other processes occurring in plants and animals maintain gametic isolation and species distinction. Species often have different mating or courtship patterns or behaviors, the breeding seasons may be distinct and even if mating does occur antigenic reactions to the sperm of other species prevent fertilization or embryo development. Hybridisation is much more common among organisms that spawn indiscriminately, like soft corals and among plants.

While it is possible to predict the genetic composition of a backcross on average, it is not possible to accurately predict the composition of a particular backcrossed individual, due to random segregation of chromosomes. In a species with two pairs of chromosomes, a twice backcrossed individual would be predicted to contain 12.5% of one species' genome (say, species A). However, it may, in fact, still be a 50% hybrid if the chromosomes from species A were lucky in two successive segregations, and meiotic crossovers happened near the telomeres. The chance of this is fairly high: ${\displaystyle \left({\frac {1}{2}}\right)^{(2\times 2)}={\frac {1}{16}}}$ (where the "two times two" comes about from two rounds of meiosis with two chromosomes); however, this probability declines markedly with chromosome number and so the actual composition of a hybrid will be increasingly closer to the predicted composition.

Hybrid species

While not common, a few animal species have been recognized as the result of hybridization. The Lonicera fly is an example of a novel animal species that resulted from natural hybridization. The American red wolf appears to be a hybrid species between gray wolf and coyote, although its taxonomic status has been a subject of controversy. The European edible frog appears to be a species, but is actually a semi-permanent hybrid between pool frogs and marsh frogs. The edible frog population is dependent on the presence of at least one of the parent species to be maintained. Evidence found in cave paintings indicates that the European bison is a descendant of the "Higgs bison", a natural hybrid between aurochs and steppe bison.

Hybrid species of plants are much more common than animals. Many of the crop species are hybrids, and hybridization appears to be an important factor in speciation in some plant groups.

Examples of hybrid animals and animal populations derived from hybrids

Mammals

• Equid hybrids
  • Mule, a cross of female horse and a male donkey
  • Hinny, a cross between a female donkey and a male horse (Mule and hinny are examples of reciprocal hybrids)
  • Zebroids
    • Zeedonk or Zonkey, a zebra/donkey cross
    • Zorse, a zebra/horse cross
    • Zony or Zetland, a zebra/pony cross ("zony" is a generic term; "zetland" is specifically a hybrid of the Shetland pony breed with a zebra)
  • hybrid ass, a cross between a donkey and an onager or Asian wild ass
• Bovid hybrids
  • Dzo, zo or yakow; a cross between a domestic cow/bull and a yak
  • Beefalo, a cross of an American bison and a domestic cow. This is a fertile breed; this along with genetic evidence has caused them to be recently reclassified into the same genus, Bos.
  • Å»ubroÅ„, a hybrid between wisent (European bison) and domestic cow
  • "Higgs bison", a naturally occurring hybrid between aurochs and steppe bison that appeared during the Pleistocene; likely gave rise to the wisent.
• Sheep-goat hybrid is the cross between a sheep and a goat, which belong to different genera.
• Ursid hybrids, such as the grizzly-polar bear hybrid, occur between black bears, brown bears, and polar bears.
• Felid hybrids
  • Savannah cat is a fertile breed developed originally from a cross between the serval (Leptailurus serval) and domestic cats (Felis catus).
  • A hybrid between a Bengal tiger and a Siberian tiger is an example of an intra-specific hybrid. It also includes the Indochinese tiger, Sumatran tiger.
  • Pumapards are the hybrid crosses between a puma and a leopard.
  • Ligers and tigons (crosses between a lion and a tiger - the difference in name due to which parent is of which species - ligers have a lion father and a tiger mother) and other Panthera hybrids such as the lijagulep. Various other wild cat crosses are known involving the lynx, bobcat, leopard, serval, etc.
    • Liligers are the hybrid cross between a male lion and a ligress.
  • Bengals are a fertile breed developed originally from a cross between the Asian leopard cat [Prionailurus bengalensis] and the domestic cat [Felis catus].
• Fertile canid hybrids occur between coyotes, wolves, dingoes, jackals and domestic dogs.
• Hybrids between black and white rhinoceroses have been recognized.
• Hybrid camel, a cross between a bactrian camel and a dromedary camel
• Cama, a cross between a camel and a llama, also an intergeneric hybrid.
• Wholphin, a fertile but very rare cross between a false killer whale and a bottlenose dolphin.
• At Chester Zoo in the United Kingdom, a cross between an African elephant (male) and an Asian elephant (female). The male calf was named Motty. He died of intestinal infection after twelve days.
• Bornean and Sumatran orangutan hybrids have occurred in captivity.

Birds

• Hybrids between spotted owls and barred owls
• Cagebird breeders sometimes breed hybrids between species of finch, such as goldfinch × canary. These birds are known as mules.
• The perlin is a peregrine falcon â€" merlin hybrid.
• Gamebird hybrids, hybrids between gamebirds and domestic fowl, including chickens, guineafowl and peafowl, interfamilial hybrids.
• Numerous macaw hybrids and lovebird hybrids are also known in aviculture.
• Red kite × black kite: five bred unintentionally at a falconry center in England. (It is reported that the black kite (the male) refused female black kites but mated with two female red kites.)
• The mulard duck, hybrid of the domestic pekin duck and domesticated muscovy ducks.
• In Australia, New Zealand and other areas where the Pacific black duck occurs, it is hybridised by the much more aggressive introduced mallard. This is a concern to wildlife authorities throughout the affected area, as it is seen as Genetic pollution of the black duck gene pool.
• Hybridisation in gulls is a reasonably frequent occurrence in the wild.

Reptiles

• Hybrid iguana, a single‐cross hybrid resulting from natural interbreeding between male marine iguanas and female land iguanas since the late 2000s.
• Crestoua, a cross between a Rhacodactylus Ciliatus (crested gecko) and a Rhacodactylus Chahoua.
• Colubrid snakes of the tribe Lampropeltini have been shown to produce fertile hybrid offspring.
• Hybridization between the endemic Cuban crocodile (Crocodylus rhombifer) and the widely distributed American crocodile (Crocodylus acutus) is causing conservation problems for the former species as a threat to its genetic integrity.
• Saltwater crocodiles (Crocodylus porosus) have mated with Siamese crocodiles (Crocodylus siamensis) in captivity producing offspring which in many cases have grown over 20 feet (6.1 metres) in length. It is likely that wild hybridization occurred historically in parts of southeast Asia.
• Many species of boas and pythons are known to produce hybrids, such as carball (a cross between a ball python and a carpet python) or a bloodball (a cross between a blood python and a ball python); however most of these only occur in captivity. Contrary to popular belief, boaâ€"python hybrids are not possible due to their differing reproductive functions. Boas only produce hybrids with other species of boas, and pythons only produce hybrids with other species of pythons.

Amphibians

• Japanese giant salamanders and Chinese giant salamanders have created hybrids that threaten the survival of Japanese giant salamanders due to the competition for similar resources in Japan.

Fish

• Blood parrot cichlid, which is probably created by crossing a red head cichlid and a Midas cichlid or red devil cichlid
• A group of about 50 hybrids between Australian blacktip shark and the larger common blacktip shark was found by Australia's East Coast in 2012. This is the only known case of hybridization in sharks.
• Silver bream and common bream commonly produce sterile hybrids.
• Tiger muskie is a sterile hybrid between northern pike and muskellunge.

Insects

• Killer bees were created during an attempt to breed a strain of bees that would produce more honey and be better adapted to tropical conditions. This was done by crossing a European honey bee and an African bee.
• The Colias eurytheme and C. philodice butterflies have retained enough genetic compatibility to produce viable hybrid offspring.

Hybrid plants

Many hybrid plants are created by humans, but natural hybrids also occur. Plant species hybridize more readily than animal species, and the resulting hybrids are more often fertile hybrids and may reproduce, though there still exist sterile hybrids and selective hybrid elimination where the offspring are less able to survive and are thus eliminated before they can reproduce. A number of plant species are the result of hybridization and polyploidy with many plant species easily cross-pollinating and producing viable seeds, the distinction between each species is often maintained by geographical isolation or differences in the flowering period. Since plants hybridize frequently without much work, they are often created by humans in order to produce improved plants. These improvements can include the production of more or improved seeds, fruits or other plant parts for consumption, or to make a plant more winter- or heat-resistant, or to improve its growth and or appearance for use in horticulture. Much work is now being done with hybrids to produce disease-resistant plants for both agricultural and horticultural crops. In many plant species, hybridization is used to produce larger and more showy flowers and new flower colors. Hybridization may be restricted to the desired parent species through the use of pollination bags.

Many plant genera and species have their origins in polyploidy. Autopolyploidy results from the sudden multiplication in the number of chromosomes in typical normal populations caused by unsuccessful separation of the chromosomes during meiosis. Tetraploids (plants with four sets of chromosomes rather than two) are common in a number of different groups of plants and over time these plants can differentiate into distinct species from the normal diploid line. In Oenothera lamarchiana the diploid species has 14 chromosomes, this species has spontaneously given rise to plants with 28 chromosomes that have been given the name Oenothera gigas. When hybrids are formed between the tetraploids and the diploid population, the resulting offspring tend to be sterile triploids, thus effectively stopping the intermixing of genes between the two groups of plants (unless the diploids, in rare cases, produce unreduced gametes).

Another form of polyploidy called allopolyploidy occurs when two different species mate and produce polyploid hybrids. Usually the typical chromosome number is doubled, and the four sets of chromosomes can pair up during meiosis, thus the polyploids can produce offspring. Usually, these offspring can mate and reproduce with each other but cannot back-cross with the parent species. Allopolyploids may be able to adapt to new habitats that neither of their parent species inhabited.

Sterility in a non-polyploid hybrid is often a result of chromosome number; if parents are of differing chromosome pair number, the offspring will have an odd number of chromosomes, leaving them unable to produce chromosomally balanced gametes. While this is undesirable in a crop such as wheat, where growing a crop which produces no seeds would be pointless, it is an attractive attribute in some fruits. Triploid bananas and watermelons are intentionally bred because they produce no seeds (and are parthenocarpic).

Heterosis

Hybrids are sometimes stronger than either parent variety, a phenomenon most common with plant hybrids, which when present is known as hybrid vigor (heterosis) or heterozygote advantage. A transgressive phenotype is a phenotype displaying more extreme characteristics than either of the parent lines. Plant breeders use several techniques to produce hybrids, including line breeding and the formation of complex hybrids. An economically important example is hybrid maize (corn), which provides a considerable seed yield advantage over open pollinated varieties. Hybrid seed dominates the commercial maize seed market in the United States, Canada and many other major maize-producing countries.

Examples of plant hybrids

The multiplication symbol × (not italicised) indicates a hybrid in the Latin binomial nomenclature. Placed before the binomial it indicates a hybrid between species from different genera (intergeneric hybrid):-

• × Fatshedera lizei, a hybrid between Hedera helix and Fatsia japonica
• × Heucherella, a hybrid genus between Heuchera and Tiarella
• × Philageria veitchii is a hybrid between Lapageria rosea and Philesia magellanica; it is more similar in appearance to the former
• Leyland cypress, [× Cupressocyparis leylandii] hybrid between Monterey cypress and Nootka cypress
• Triticale, [× Triticosecale] a wheatâ€"rye hybrid
• × Urceocharis, a hybrid between Eucharis and Urceolina

Interspecific plant hybrids include:

• Dianthus × allwoodii (Dianthus caryophyllus × Dianthus plumarius)
• Limequat Citrus × floridana, key lime Citrus aurantiifolia and kumquat Citrus japonica hybrid
• Loganberry Rubus × loganobaccus, a hybrid between raspberry Rubus idaeus and blackberry Rubus ursinus
• London plane (Platanus orientalis × Platanus occidentalis), thus forming Platanus × acerifolia
• Magnolia × alba (Magnolia champaca × Magnolia montana)
• Peppermint, a hybrid between spearmint and water mint
• Quercus × warei (Quercus robur × Quercus bicolor) 'Nadler' (marketed in the United States under the trade name Kindred Spirit hybrid oak)
• Tangelo, a hybrid of a Mandarin orange and a pomelo which may have been developed in Asia about 3,500 years ago
• Wheat; most modern and ancient wheat breeds are themselves hybrids. Bread wheat is a hexaploid hybrid of three wild grasses; durum (pasta) wheat is a tetraploid hybrid of two wild grasses
• Grapefruit, hybrid between a pomelo and the Jamaican sweet orange

Some natural hybrids:

• Iris albicans, a sterile hybrid which spreads by rhizome division
• Evening primrose, a flower which was the subject of famous experiments by Hugo de Vries on polyploidy and diploidy

Hybrids in nature

Hybridization between two closely related species is actually a common occurrence in nature but is also being greatly influenced by anthropogenic changes as well. Hybridization is a naturally occurring genetic process where individuals from two genetically distinct populations mate. It can occur both intraspecifically, between different distinct populations within the same species, and interspecifically, between two different species. Hybrids can be either sterile/not viable or viable/fertile. This affects the impact which the hybrid will have on its and other populations that it interacts with. Many hybrid zones are known where the ranges of two species meet, and hybrids are continually produced in great numbers. These hybrid zones are useful as biological model systems for studying the mechanisms of speciation (Hybrid speciation). Recently DNA analysis of a bear shot by a hunter in the North West Territories confirmed the existence of naturally-occurring and fertile grizzlyâ€"polar bear hybrids.

Anthropogenic hybridization

Changes to the environment caused by humans, such as fragmentation and Introduced species, are becoming more widespread. This increases the challenges in managing certain populations that are experiencing introgression, and is a focus of conservation genetics.

Introduced species and habitat fragmentation

Humans have introduced species worldwide to environments for a long time, both intentionally such as establishing a population to be used as a biological control, and unintentionally such as accidental escapes of individuals out of agriculture. This causes drastic global effects on various populations, including through hybridization.

When habitats become broken apart, one of two things can occur, genetically speaking. The first is that populations that were once connected can be cut off from one another, preventing their genes from interacting. Occasionally, this will result in a population of one species breeding with a population of another species as a means of surviving such as the case with the red wolves. Their population numbers being so small, they needed another means of survival. Habitat fragmentation also led to the influx of generalist species into areas where they would not have been, leading to competition and in some cases interbreeding/incorporation of a population into another. In this way, habitat fragmentation is essentially an indirect method of introducing species to an area.

The hybridization continuum

There is a kind of continuum with three semi-distinct categories dealing with anthropogenic hybridization: hybridization without Introgression, hybridization with widespread introgression, and essentially a Hybrid swarm. Depending on where a population falls along this continuum, the management plans for that population will change. Hybridization is currently an area of great discussion within Wildlife management and habitat management. Global climate change is creating other changes such as difference in population distributions which are indirect causes for an increase in anthropogenic hybridization.

Consequences

Hybridization can be a less discussed way toward extinction than within detection of where a population lies along the hybrid continuum. The dispute of hybridization is how to manage the resulting hybrids. When a population experiences hybridization with substantial introgression, there still exists parent types of each set of individuals. When a complete hybrid swarm is created, all the individuals are hybrids.

Management of hybrids

Conservationists disagree on when is the proper time to give up on a population that is becoming a hybrid swarm or to try and save the still existing pure individuals. Once it becomes a complete mixture, we should look to conserve those hybrids to avoid their loss. Most leave it as a case-by-case basis, depending on detecting of hybrids within the group. It is nearly impossible to regulate hybridization via policy because hybridization can occur beneficially when it occurs "naturally" and there is the matter of protecting those previously mentioned hybrid swarms because if they are the only remaining evidence of prior species, they need to be conserved as well.

Expression of parental traits in hybrids

When two distinct types of organisms breed with each other, the resulting hybrids typically have intermediate traits (e.g., one parent has red flowers, the other has white, and the hybrid, pink flowers). Commonly, hybrids also combine traits seen only separately in one parent or the other (e.g., a bird hybrid might combine the yellow head of one parent with the orange belly of the other).

In a hybrid, any trait that falls outside the range of parental variation is termed heterotic. Heterotic hybrids do have new traits, that is, they are not intermediate. Positive heterosis produces more robust hybrids, they might be stronger or bigger; while the term negative heterosis refers to weaker or smaller hybrids. Heterosis is common in both animal and plant hybrids. For example, hybrids between a lion and a tigress ("ligers") are much larger than either of the two progenitors, while a tigon (lioness × tiger) is smaller. Also the hybrids between the common pheasant (Phasianus colchicus) and domestic fowl (Gallus gallus) are larger than either of their parents, as are those produced between the common pheasant and hen golden pheasant (Chrysolophus pictus). Spurs are absent in hybrids of the former type, although present in both parents.

Genetic mixing and extinction

Regionally developed ecotypes can be threatened with extinction when new alleles or genes are introduced that alter that ecotype. This is sometimes called genetic mixing. Hybridization and introgression of new genetic material can lead to the replacement of local genotypes if the hybrids are more fit and have breeding advantages over the indigenous ecotype or species. These hybridization events can result from the introduction of non-native genotypes by humans or through habitat modification, bringing previously isolated species into contact. Genetic mixing can be especially detrimental for rare species in isolated habitats, ultimately affecting the population to such a degree that none of the originally genetically distinct population remains.

Effect on biodiversity and food security

In agriculture and animal husbandry, the Green Revolution's use of conventional hybridization increased yields by breeding "high-yielding varieties". The replacement of locally indigenous breeds, compounded with unintentional cross-pollination and crossbreeding (genetic mixing), has reduced the gene pools of various wild and indigenous breeds resulting in the loss of genetic diversity. Since the indigenous breeds are often well-adapted to local extremes in climate and have immunity to local pathogens, this can be a significant genetic erosion of the gene pool for future breeding. Therefore, commercial plant geneticists strive to breed "widely adapted" cultivars to counteract this tendency.

Limiting factors

A number of conditions exist that limit the success of hybridization; the most obvious is great genetic diversity between most species. But in animals and plants that are more closely related, hybridization barriers can include morphological differences, differing times of fertility, mating behaviors and cues, physiological rejection of sperm cells or the developing embryo.

In plants, barriers to hybridization include blooming period differences, different pollinator vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and structural differences of the chromosomes.

Mythical, legendary and religious hybrids

Ancient folktales often contain mythological creatures, sometimes described as hybrids (e.g., hippogriff as the offspring of a griffin and a horse, and Minotaur, the offspring of Pasiphaë and a white bull). More often they are kind of chimera, i.e., a composite of the physical attributes of two or more kinds of animals, mythical beasts, and often humans, with no suggestion that they are the result of interbreeding, e.g., harpies, mermaids, and centaurs.

In the Bible, the Old Testament contains several passages which talk about a first generation of hybrid giants who were known as the Nephilim. The Book of Genesis (6:4) states that "the sons of God went to the daughters of humans and had children by them". As a result, the offspring were born as hybrid giants who became mighty heroes of old and legendary famous figures of ancient times. In addition, the Book of Numbers (13:33) says that the descendants of Anak came from the Nephilim, whose bodies looked exactly like men, but with an enormous height. According to the apocryphal Book of Enoch the Nephilim were wicked sons of fallen angels who had lusted with attractive women.

See also

References

External links

• Artificial Hybridisation â€" Artificial Hybridisation in orchids
• Domestic Fowl Hybrids
• Hybrid Mammals
• Hybridisation in animals Evolution Revolution: Two Species Become One, Study Says (nationalgeographic.com)
• Hybrids of wildcats with domestic cats
• Scientists Create Butterfly Hybrid â€" Creation of new species through hybridization was thought to be common only in plants, and rare in animals.
• What is a human admixed embryo?
• Video of Mirror carp & Gold fish spawning at a fishing venue in France