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90-426: The tigon is a hybrid offspring of a male tiger ( Panthera tigris ) and a female lion , or lioness ( Panthera leo ). They exhibit visible characteristics from both parents: they can have both spots from the mother (lions carry genes for spots – lion cubs are spotted and some adults retain faint markings) and stripes from the father. Any mane that a male tigon may have will appear shorter and less noticeable than

180-475: A synthetic population . In horticulture , the term stable hybrid is used to describe an annual plant that, if grown and bred in a small monoculture free of external pollen (e.g., an air-filtered greenhouse) produces offspring that are "true to type" with respect to phenotype; i.e., a true-breeding organism. Hybridization can occur in the hybrid zones where the geographical ranges of species, subspecies, or distinct genetic lineages overlap. For example,

270-463: A bactrian camel and a dromedary . There are many examples of felid hybrids , including the liger . The oldest-known animal hybrid bred by humans is the kunga equid hybrid produced as a draft animal and status symbol 4,500 years ago in Umm el-Marra , present-day Syria . The first known instance of hybrid speciation in marine mammals was discovered in 2014. The clymene dolphin ( Stenella clymene )

360-588: A 15-year-old hybrid between a lion and a tiger was successfully mated at the Munich Hellabrunn Zoo . The female cub was then raised to adulthood. Like the liger, male tigons are sterile while the females are fertile. At the Alipore Zoo in India , a tigoness named Rudhrani, born in 1971, was successfully mated to a male Asiatic lion named Debabrata. The rare, second-generation hybrid was called

450-564: A bear shot by a hunter in the Northwest Territories confirmed the existence of naturally occurring and fertile grizzly–polar bear hybrids . Hybridization between reproductively isolated species often results in hybrid offspring with lower fitness than either parental. However, hybrids are not, as might be expected, always intermediate between their parents (as if there were blending inheritance), but are sometimes stronger or perform better than either parental lineage or variety,

540-672: A class of non-coding small RNAs which repress the translation of messenger RNAs (mRNAs) or cause degradation of mRNAs. In hybrid plants, most miRNAs have non-additive expression (it might be higher or lower than the levels in the parents). This suggests that the small RNAs are involved in the growth, vigor and adaptation of hybrids. 'Heterosis without hybridity' effects on plant size have been demonstrated in genetically isogenic F1 triploid (autopolyploid) plants, where paternal genome excess F1 triploids display positive heterosis, whereas maternal genome excess F1s display negative heterosis effects. Such findings demonstrate that heterosis effects, with

630-502: 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', and nonstiff stalk. Hybrid rice sees cultivation in many countries, including China, India, Vietnam, and

720-515: 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. In a hybrid, any trait that falls outside the range of parental variation (and is thus not simply intermediate between its parents) is considered heterotic. Positive heterosis produces more robust hybrids, they might be stronger or bigger; while

810-509: A cross between a female horse and a male donkey, and the hinny, a cross between a female donkey and a male horse. Pairs of complementary types like the mule and hinny are called reciprocal hybrids. Polar bears and brown bears are another case of a hybridizing species pairs, and introgression among non-sister species of bears appears to have shaped the Ursidae family tree. Among many other mammal crosses are hybrid camels , crosses between

900-509: A degree that none of the originally genetically distinct population remains. 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

990-401: A detailed study of purebred Cocker Spaniels, purebred Basenjis, and hybrids between them. They found that hybrids ran faster than either parent, perhaps due to heterosis. Other characteristics, such as basal heart rate, did not show any heterosis—the dog's basal heart rate was close to the average of its parents—perhaps due to the additive effects of multiple genes. Sometimes people working on

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1080-669: A dog-breeding program find no useful heterosis. All this said, studies do not provide definitive proof of hybrid vigor in dogs. This is largely due to the unknown heritage of most mixed breed dogs used. Results vary wildly, with some studies showing benefit and others finding the mixed breed dogs to be more prone to genetic conditions. In 2014, a study undertaken by the Centre for Integrative Ecology at Deakin University in Geelong, Victoria, concluded that intraspecific hybrids between

1170-602: A donkey as the father. A variety of mechanisms limit the success of hybridization, including the large genetic difference between most species. Barriers include morphological differences, differing times of fertility, mating behaviors and cues, and physiological rejection of sperm cells or the developing embryo. Some act before fertilization; others after it. In plants, some 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

1260-523: A genome dosage-dependent epigenetic basis, can be generated in F1 offspring that are genetically isogenic (i.e. harbour no heterozygosity). It has been shown 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 or methylation of specific amino acids in histone H3 ,

1350-548: A group of about 50 natural hybrids between Australian blacktip shark and the larger common blacktip shark was found by Australia's eastern coast in 2012. Russian sturgeon and American paddlefish were hybridized in captivity when sperm from the paddlefish and eggs from the sturgeon were combined, unexpectedly resulting in viable offspring. This hybrid is called a sturddlefish . The two genera Asymmetron and Branchiostoma are able to produce viable hybrid offspring, even if none have lived into adulthood so far, despite

1440-435: A hybrid inherits traits from its parents that are not fully compatible, fitness can be reduced. This is a form of outbreeding depression , the effects of which are similar to inbreeding depression. Since the early 1900s, 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. When

1530-500: A hybrid is interpreted differently in animal and plant breeding, where there is interest in the individual parentage. In genetics , attention is focused on the numbers of chromosomes . In taxonomy, a key question is how closely related the parent species are. Species are reproductively isolated by strong barriers to hybridization, which include genetic and morphological differences, differing times of fertility, mating behaviors and cues, and physiological rejection of sperm cells or

1620-543: 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

1710-545: A lion's mane and is closer in type to the ruff of a male tiger. Tigons do not exceed the size of their parent species because they inherit growth-inhibitory genes from both parents, but they do not exhibit any kind of dwarfism or miniaturization; they often weigh around 180 kg (400 lb). It is distinct from the liger , which is a hybrid of a male lion and a female tiger, often weighing from 320 kg (710 lb) to 550 kg (1,210 lb). Ligers and tigons were long thought to be sterile ; in 1943, however,

1800-527: A litigon. Rudhrani produced seven litigons in her lifetime. Some of these reached impressive sizes - a litigon named Cubanacan weighed at least 363 kg (800 lb), stood 1.32 m (4 ft 4 in) at the shoulder, and was 3.5 m (11 ft) in total length. Reports also exist of the similar titigon, resulting from the cross between a female tigon and a male tiger. Titigons resemble golden tigers , but with less contrast in their markings. A tigoness born in 1978, named Noelle, shared an enclosure in

1890-439: A long time, both intentionally for purposes such as biological control , and unintentionally, as with accidental escapes of individuals. Introductions can drastically affect populations, including through hybridization. There is a kind of continuum with three semi-distinct categories dealing with anthropogenic hybridization: hybridization without introgression, hybridization with widespread introgression (backcrossing with one of

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1980-417: A maintainer line, and a restorer line. The second generation was widely adopted in the 1990s. Instead of a CMS line, it uses an environment-sensitive genic male sterile line (EGMS), which can have its sterility reversed based on light or temperature. This removes the need for a maintainer, making the hybridization and breeding process more efficient (albeit still high-maintenance). Second generation lines show

2070-498: A mate among the species that raised it, instead of the species of its biological parents. Cagebird breeders sometimes breed bird hybrids known as mules between species of finch , such as goldfinch × canary . Among amphibians, Japanese giant salamanders and Chinese giant salamanders have created hybrids that threaten the survival of Japanese giant salamanders because of competition for similar resources in Japan. Among fish,

2160-455: A more psychoactive cannabis . 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

2250-432: A phenomenon called heterosis, hybrid vigour, or heterozygote advantage . This is most common with plant hybrids. A transgressive phenotype is a phenotype that displays 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

2340-531: A population is small or inbred, it tends to lose genetic diversity. Inbreeding depression is the loss of fitness due to loss of genetic diversity. Inbred strains tend to be homozygous for recessive alleles that are mildly harmful (or produce a trait that is undesirable from the standpoint of the breeder). Heterosis or hybrid vigor, on the other hand, is the tendency of outbred strains to exceed both inbred parents in fitness. Selective breeding of plants and animals, including hybridization, began long before there

2430-432: A protein closely associated with DNA, which can either activate or repress associated genes. One example of where particular genes may be important in vertebrate animals for heterosis is the major histocompatibility complex (MHC). 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

2520-405: A species, such as between different breeds . Single cross hybrids result from the cross between two true-breeding organisms which produces an F1 hybrid (first filial generation). 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,

2610-450: A yield increase of 5-10% over first generation lines. The third and current generation uses a nuclear male sterile line (NMS). Third generation lines have a recessive sterility gene, and their cultivation is more lenient towards maintainer lines and environmental conditions. Additionally, transgenes are only present in the maintainer, so hybrid plants can benefit from hybrid vigor without requiring special oversight. The concept of heterosis

2700-516: Is a plan to shift some lions from their current home of the Gir Forest to Kuno Wildlife Sanctuary , which has some tigers, but it has not been implemented as of December 2017, perhaps due to political reasons, as the Gujarat state government does not want any other state to have lions in the forests. Hybrid (biology) In biology , a hybrid is the offspring resulting from combining

2790-577: Is a hybrid of two Atlantic species, the spinner and striped dolphins . In 2019, scientists confirmed that a skull found 30 years earlier was a hybrid between the beluga whale and narwhal , dubbed the narluga . Hybridization between species is common in birds. Hybrid birds are purposefully bred by humans, but hybridization is also common in the wild. Waterfowl have a particularly high incidence of hybridization, with at least 60% of species known to produce hybrids with another species. Among ducks , mallards widely hybridize with many other species, and

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2880-407: Is a hybridization test widely used in genetics to determine whether two separately isolated mutants that have the same (or similar) phenotype are defective in the same gene or in different genes (see Complementation (genetics) article). If a hybrid organism containing the genomes of two different mutant parental organisms displays a wild type phenotype, it is ordinarily considered that

2970-571: Is a particularly common mechanism for speciation in plants, and is now known to be fundamental to the evolutionary history of plants. Plants frequently form polyploids , individuals with more than two copies of each chromosome. Whole genome doubling has occurred repeatedly in plant evolution. When two plant species hybridize, the hybrid may double its chromosome count by incorporating the entire nuclear genome of both parents, resulting in offspring that are reproductively incompatible with either parent because of different chromosome counts. Human impact on

3060-399: Is a self-pollinating species, it requires the use of male-sterile lines to generate hybrids from separate lineages. The most common way of achieving this is using lines with genetic male-sterility, as manual emasculation is not optimal for large-scale hybridization. The first generation of hybrid rice was developed in the 1970s. It relies on three lines: a cytoplasmic male sterile (CMS) line,

3150-870: Is also applied in the production of commercial livestock . In cattle, crosses between Black Angus and Hereford produce a cross known as a " Black Baldy ". In swine , "blue butts" are produced by the cross of Hampshire and Yorkshire. Other, more exotic hybrids (two different species, so genetically more dissimilar), such as " beefalo " which are hybrids of cattle and bison, 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,

3240-659: Is also more occasionally done in the livestock and pet trades; some well-known wild × domestic hybrids are beefalo and wolfdogs . Human selective breeding of domesticated animals and plants has also resulted in the development of distinct breeds (usually called cultivars in reference to plants); crossbreeds between them (without any wild stock ) are sometimes also imprecisely referred to as "hybrids". Hybrid humans existed in prehistory. For example, Neanderthals and anatomically modern humans are thought to have interbred as recently as 40,000 years ago. Mythological hybrids appear in human culture in forms as diverse as

3330-451: Is always sterile worker ants (and because ants are haplodiploid , unfertilized eggs become males). Without mating with males of the other species, the queens are unable to produce workers, and will fail to establish a colony of their own. Plant species hybridize more readily than animal species, and the resulting hybrids are fertile more often. Many plant species are the result of hybridization, combined with polyploidy , which duplicates

3420-552: Is derived from Latin hybrida , used for crosses such as of a tame sow and a wild boar. 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. Conspicuous hybrids are popularly named with portmanteau words , starting in the 1920s with the breeding of tiger–lion hybrids ( liger and tigon ). From the point of view of animal and plant breeders, there are several kinds of hybrid formed from crosses within

3510-568: 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

3600-427: Is that hybrid individuals can form a "bridge" transmitting potentially helpful genes from one species to another when the hybrid backcrosses with one of its parent species, a process called introgression . Hybrids can also cause speciation , either because the hybrids are genetically incompatible with their parents and not each other, or because the hybrids occupy a different niche than either parent. Hybridization

3690-412: Is the main cause for the fitness advantages of heterosis, then there should be an over-expression of certain genes in the heterozygous offspring compared to the homozygous parents. On the other hand, if dominance is the cause, fewer genes should be under-expressed in the heterozygous offspring compared to the parents. Furthermore, for any given gene, the expression should be comparable to the one observed in

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3780-515: 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 a population becomes a complete mixture, the goal becomes to conserve those hybrids to avoid their loss. Conservationists treat each case on its merits, depending on detecting hybrids within the population. It is nearly impossible to formulate a uniform hybridization policy, because hybridization can occur beneficially when it occurs "naturally", and when hybrid swarms are

3870-635: The Flynn effect , the steady rise in IQ test scores around the world during the 20th century, though a review of nine studies found that there is no evidence to suggest inbreeding has an effect on IQ. The term heterosis often causes confusion and even controversy, particularly in selective breeding of domestic animals , because it is sometimes (incorrectly) claimed that all crossbred plants and animals are "genetically superior" to their parents, due to heterosis, . but two problems exist with this claim: An example of

3960-653: The Minotaur , blends of animals, humans and mythical beasts such as centaurs and sphinxes , and the Nephilim of the Biblical apocrypha described as the wicked sons of fallen angels and attractive women. Hybridization between species plays an important role in evolution, though there is much debate about its significance. Roughly 25% of plants and 10% of animals are known to form hybrids with at least one other species. One example of an adaptive benefit to hybridization

4050-472: The sea urchin Strongylocentrotus purpuratus (female) and the sand dollar Dendraster excentricus (male). When two distinct types of organisms breed with each other, the resulting hybrids typically have intermediate traits (e.g., one plant 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

4140-438: The "suture region". It is at these regions that the subspecies were formed. Other hybrid zones have formed between described species of plants and animals. From the point of view of genetics, several different kinds of hybrid can be distinguished. A genetic hybrid carries two different alleles of the same gene , where for instance one allele may code for a lighter coat colour than the other. A structural hybrid results from

4230-638: The 1980s, as experimental evidence has mounted, the dominance theory has made a comeback. Crow wrote: The current view ... is that the dominance hypothesis is the major explanation of inbreeding decline and [of] the high yield of hybrids. There is little statistical evidence for contributions from overdominance and epistasis . But whether the best hybrids are getting an extra boost from overdominance or favorable epistatic contributions remains an open question. An epigenetic contribution to heterosis has been established in plants, and it has also been reported in animals. MicroRNAs (miRNAs), discovered in 1993, are

4320-615: 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 at a marketable carcass weight 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. In 2013, a study found that mixed breeds live on average 1.2 years longer than pure breeds. John Scott and John L. Fuller performed

4410-458: 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 (i.e., there are four unrelated grandparents). 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. Top cross (or "topcross") hybrids result from

4500-571: The Philippines. Compared to inbred lines, hybrids produce approximately 20% greater yield, and comprise 45% of rice planting area in China. 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 around 15 tons per hectare. In India also, several varieties have shown high vigor, including 'RH-10' and 'Suruchi 5401'. Since rice

4590-485: The Shambala Preserve with a male Siberian tiger called Anton, due to the keepers' belief that she was sterile. In 1983 Noelle produced a titigon named Nathaniel. As Nathaniel was three-quarters tiger, he had darker stripes than Noelle and vocalized more like a tiger, rather than with the mix of sounds used by his mother. Being only about quarter-lion, Nathaniel did not grow a mane. Nathaniel died of cancer at

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4680-612: The age of nine years old. Noelle also developed a severe cancer that killed her not long after she was diagnosed. As with the liger, the tigon is found only in captivity, because the habitats of the tiger and lion do not overlap. In the past, however, the Asiatic lion did coexist with the Bengal tiger in the wilderness of India, besides occurring in countries where the Caspian tiger had been, such as Iran and Turkey . In India, there

4770-417: The ambiguous value judgements imposed 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

4860-524: The butterfly Limenitis arthemis has two major subspecies in North America, L. a. arthemis (the white admiral) and L. a. astyanax (the red-spotted purple). The white admiral has a bright, white band on its wings, while the red-spotted purple has cooler blue-green shades. Hybridization occurs between a narrow area across New England, southern Ontario, and the Great Lakes,

4950-584: The cause, though, of autoimmune diseases. 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

5040-503: The chance that any particular pathogen will be recognised, and means that more antigenic proteins on any pathogen are likely to be recognised, giving a greater range of T-cell activation, so a greater response. This also means that the immunity acquired to the pathogen is against a greater range of antigens, meaning that the pathogen must mutate more before immunity is lost. Thus, hybrids are less likely to succumb to pathogenic disease and are more capable of fighting off infection. This may be

5130-426: The chromosomes. A few animal species are the result of hybridization. The Lonicera fly is a natural hybrid. The American red wolf appears to be a hybrid of the gray wolf and the coyote , although its taxonomic status has been a subject of controversy. The European edible frog is a semi-permanent hybrid between pool frogs and marsh frogs ; its population requires the continued presence of at least one of

5220-547: The chromosomes. Chromosome duplication allows orderly meiosis and so viable seed can be produced. Plant hybrids are generally given names that include an "×" (not in italics), such as Platanus × hispanica for the London plane, a natural hybrid of P. orientalis (oriental plane) and P. occidentalis (American sycamore). The parent's names may be kept in their entirety, as seen in Prunus persica × Prunus americana , with

5310-481: The common pheasant and hen golden pheasant ( Chrysolophus pictus ). Spurs are absent in hybrids of the former type, although present in both parents. Hybridization is greatly influenced by human impact on the environment, through effects such as habitat fragmentation and species introductions. Such impacts make it difficult to conserve the genetics of populations undergoing introgressive hybridization . Humans have introduced species worldwide to environments for

5400-400: The crossing of a top quality or pure-bred male and a lower quality female, intended to improve the quality of the offspring, on average. Population hybrids result from the crossing of plants or animals in one population with those of another population. These include interspecific hybrids or crosses between different breeds. In biology, the result of crossing of two populations is called

5490-444: The developing embryo . Some act before fertilization and others after it. Similar barriers exist in plants, with differences in flowering times, pollen vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and the structure of the chromosomes. A few animal species and many plant species, however, are the result of hybrid speciation , including important crop plants such as wheat , where

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5580-407: The diverse Heliconius butterflies , but that is disputed. The two closely related harvester ant species Pogonomyrmex barbatus and Pogonomyrmex rugosus have evolved to depend on hybridization. When a queen fertilizes her eggs with sperm from males of her own species, the offspring is always new queens. And when she fertilizes the eggs with sperm from males of the other species, the offspring

5670-413: The doubling of chromosome sets, causing immediate genetic isolation. Hybridization may be important in speciation in some plant groups. However, homoploid hybrid speciation (not increasing the number of sets of chromosomes) may be rare: by 1997, only eight natural examples had been fully described. Experimental studies suggest that hybridization offers a rapid route to speciation, a prediction confirmed by

5760-611: The environment has resulted in an increase in the interbreeding between regional species, and the proliferation of introduced species worldwide has also resulted in an increase in hybridization. This has been referred to as genetic pollution out of concern that it may threaten many species with extinction. Similarly, genetic erosion from monoculture in crop plants may be damaging the gene pools of many species for future breeding. The conservation impacts of hybridization between species are highly debated. While hybridization could potentially threaten rare species or lineages by "swamping"

5850-429: The fact that early generation hybrids and ancient hybrid species have matching genomes, meaning that once hybridization has occurred, the new hybrid genome can remain stable. 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. Recently DNA analysis of

5940-496: The female parent's name given first, or if not known, the parent's names given alphabetically. Heterosis 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. The physiological vigor of an organism as manifested in its rapidity of growth, its height and general robustness,

6030-628: The field of genetics . Population geneticist James Crow (1916–2012) believed, in his younger days, that overdominance was a major contributor to hybrid vigor. In 1998 he published a retrospective review of the developing science. According to Crow, the demonstration of several cases of heterozygote advantage in Drosophila and other organisms first caused great enthusiasm for the overdominance theory among scientists studying plant hybridization. But overdominance implies that yields on an inbred strain should decrease as inbred strains are selected for

6120-416: The fitter of the two parents. 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 . Which of the two mechanisms are the "main" reason for heterosis has been a scientific controversy in

6210-565: The fusion of gametes that have differing structure in at least one chromosome, as a result of structural abnormalities . A numerical hybrid results from the fusion of gametes having different haploid numbers of chromosomes . A permanent hybrid results when only the heterozygous genotype occurs, as in Oenothera lamarckiana , because all homozygous combinations are lethal. In the early history of genetics, Hugo de Vries supposed these were caused by mutation . Genetic complementation

6300-438: 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 are more similar in smaller, more closely related populations. Breeding between more genetically distant individuals decreases the chance of inheriting two alleles that are the same or similar, allowing a more diverse range of peptides to be presented. This, therefore, increases

6390-507: The genetic relationships between ducks are further complicated by the widespread gene flow between wild and domestic mallards. One of the most common interspecific hybrids in geese occurs between Greylag and Canada geese ( Anser anser x Branta canadensis ). One potential mechanism for the occurrence of hybrids in these geese is interspecific nest parasitism , where an egg is laid in the nest of another species to be raised by non-biological parents. The chick imprints upon and eventually seeks

6480-630: The genetically "pure" individuals with hybrids, hybridization could also save a rare lineage from extinction by introducing genetic diversity. It has been proposed that hybridization could be a useful tool to conserve biodiversity by allowing organisms to adapt, and that efforts to preserve the separateness of a "pure" lineage could harm conservation by lowering the organisms' genetic diversity and adaptive potential, particularly in species with low populations. While endangered species are often protected by law, hybrids are often excluded from protection, resulting in challenges to conservation. The term hybrid

6570-401: 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

6660-400: 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. Familiar examples of equid hybrids are the mule,

6750-452: The number of chromosomes has been doubled. A form of often intentional human-mediated hybridization is the crossing of wild and domesticated species. This is common in both traditional horticulture and modern agriculture ; many commercially useful fruits, flowers, garden herbs, and trees have been produced by hybridization. One such flower, Oenothera lamarckiana , was central to early genetics research into mutationism and polyploidy. It

6840-420: The only remaining evidence of prior species, they need to be conserved as well. 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, which can happen in natural and hybrid populations, of new genetic material can lead to the replacement of local genotypes if

6930-438: The other (e.g., a bird hybrid might combine the yellow head of one parent with the orange belly of the other). Interspecific hybrids are bred by mating individuals from two species, normally from within the same genus. The offspring display traits and characteristics of both parents, but are often sterile , preventing gene flow between the species. Sterility is often attributed to the different number of chromosomes between

7020-573: The parent species), and hybrid swarms (highly variable populations with much interbreeding as well as backcrossing with the parent species). 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. Conservationists disagree on when

7110-537: The parent species. Cave paintings indicate that the European bison is a natural hybrid of the aurochs and the steppe bison . Plant hybridization is more commonplace compared to animal hybridization. Many crop species are hybrids, including notably the polyploid wheats : some have four sets of chromosomes (tetraploid) or six (hexaploid), while other wheat species have (like most eukaryotic organisms) two sets ( diploid ), so hybridization events likely involved

7200-506: The parents' common ancestor living tens of millions of years ago. Among insects, so-called killer bees were accidentally created during an attempt to breed a strain of bees that would both produce more honey and be better adapted to tropical conditions. It 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 speciation may have produced

7290-443: The performance of their hybrid crosses, as the proportion of harmful recessives in the inbred population rises. Over the years, experimentation in plant genetics has proven that the reverse occurs, that yields increase in both the inbred strains and the hybrids, suggesting that dominance alone may be adequate to explain the superior yield of hybrids. Only a few conclusive cases of overdominance have been reported in all of genetics. Since

7380-637: The point of view of taxonomy , hybrids differ according to their parentage. Hybrids between different subspecies (such as between the dog and Eurasian wolf ) are called intra-specific hybrids. Interspecific hybrids are the offspring from interspecies mating ; these sometimes result in hybrid speciation. Intergeneric hybrids result from matings between different genera, such as between sheep and goats . Interfamilial hybrids, such as between chickens and guineafowl or pheasants , are reliably described but extremely rare. Interordinal hybrids (between different orders) are few, but have been engineered between

7470-580: The qualities of two organisms of different varieties , subspecies , species or genera through sexual reproduction . Generally, it means that each cell has genetic material from two different organisms, whereas an individual where some cells are derived from a different organism is called a chimera . Hybrids are not always intermediates between their parents such as in blending inheritance (a now discredited theory in modern genetics by particulate inheritance ), but can show hybrid vigor , sometimes growing larger or taller than either parent. The concept of

7560-405: The subspecies Platycercus elegans flaveolus and P. e. elegans of the crimson rosella ( P. elegans ) were more likely to fight off diseases than their pure counterparts. Human beings are all extremely genetically similar to one another. Michael Mingroni has proposed heterosis, in the form of hybrid vigor associated with historical reductions of the levels of inbreeding, as an explanation of

7650-470: 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 " tigons " (lioness × tiger) are smaller. Similarly, 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

7740-400: The two parental mutant organisms are defective in different genes. If the hybrid organism displays a distinctly mutant phenotype, the two mutant parental organisms are considered to be defective in the same gene. However, in some cases the hybrid organism may display a phenotype that is only weakly (or partially) wild-type, and this may reflect intragenic (interallelic) complementation. From

7830-462: The two species. For example, donkeys have 62 chromosomes , horses have 64 chromosomes, and mules or 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

7920-489: 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

8010-579: 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

8100-423: Was an understanding of underlying scientific principles. In the early 20th century, after Mendel's laws came to be understood and accepted, geneticists undertook to explain the superior vigor of many plant hybrids. Two competing hypotheses, which are not mutually exclusive, were developed: Dominance and overdominance have different consequences for the gene expression profile of the individuals. If overdominance

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