Introgression

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A phylogenetic model of introgressive hybridization; the hybrid zone of the two species' lineages is shown in blue, with each horizontal line representing an individual introgressive event. Hybrid PNGT.png
A phylogenetic model of introgressive hybridization; the hybrid zone of the two species' lineages is shown in blue, with each horizontal line representing an individual introgressive event.

Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introgression is a long-term process, even when artificial; it may take many hybrid generations before significant backcrossing occurs. This process is distinct from most forms of gene flow in that it occurs between two populations of different species, rather than two populations of the same species.

Contents

Introgression also differs from simple hybridization. Simple hybridization results in a relatively even mixture; gene and allele frequencies in the first generation will be a uniform mix of two parental species, such as that observed in mules. Introgression, on the other hand, results in a complex, highly variable mixture of genes, and may only involve a minimal percentage of the donor genome.

Definition

Introgression or introgressive hybridization is the incorporation (usually via hybridization and backcrossing) of novel genes or alleles from one taxon into the gene pool of a second, distinct taxon. [1] [2] [3] [4] This introgression is considered 'adaptive' if the genetic transfer results in an overall increase in the recipient taxon's fitness. [5]

Ancient introgression events can leave traces of extinct species in present-day genomes, a phenomenon known as ghost introgression. [6]

Source of variation

Introgression is an important source of genetic variation in natural populations and may contribute to adaptation and even adaptive radiation. [7] It can occur across hybrid zones due to chance, selection or hybrid zone movement. [8] There is evidence that introgression is a ubiquitous phenomenon in plants and animals, [9] [10] including humans, [11] in which it may have introduced the microcephalin D allele. [12]

It has been proposed that, historically, introgression with wild animals is a large contributor to the wide range of diversity found in domestic animals, rather than multiple independent domestication events. [13]

Introgressive hybridization has also been shown to be important in the evolution of domesticated crop species, possibly providing genes that help in their expansion into different environments. A genomic study from New York University Abu Dhabi Center for Genomics and Systems Biology showed that domesticated date palm varieties from North Africa show introgressive hybridization of between 5–18% of its genome from the wild Cretan palm Phoenix theophrasti into Middle East date palms P. dactylifera. This process is also similar to the evolution of apples by hybridization of Central Asian apples with the European crabapple. [14] It has also been shown that indica rice arose when Chinese japonica rice arrived in India about ~4,500 years ago and hybridized with an undomesticated proto-indica or wild O. nivara, and transferred key domestication genes from japonica to indica. [15]

Examples

Humans

There is strong evidence for the introgression of Neanderthal genes [16] and Denisovan genes [17] into parts of the modern human gene pool.

Birds

The Mallard duck is possibly the world's most capable bird to hybridise with other duck species, often to the point of the loss of genetic identity of these species. For example, feral mallard populations have significantly reduced wild populations of the Pacific black duck in New Zealand and Australia through cross-breeding.[ citation needed ]

Butterflies

One important example of introgression has been observed in studies of mimicry in the butterfly genus Heliconius. [18] This genus includes 43 species and many races with different color patterns. Congeners exhibiting overlapping distributions show similar color patterns. The subspecies H. melpomene amaryllis and H. melpomene timareta ssp. nov. overlap in distribution.

Using the ABBA/BABA test, some researchers have observed that there is about 2% to 5% introgression between the pair of subspecies. Importantly, the introgression is not random. The researchers saw significant introgression in chromosomes 15 and 18, where important mimicry loci are found (loci B/D and N/Yb). They compared both subspecies with H. melpomene agalope, which is a subspecies near H. melpomene amaryllis in entire genome trees. The result of the analysis was that there is no relation between those two species and H. melpomene agalope in the loci B/D and N/Yb. Moreover, they performed the same analysis with two other species with overlapping distributions, H. timareta florencia and H. melpomene agalope. They demonstrated introgression between the two taxa, especially in the loci B/D and N/Yb.

Finally, they concluded their experiments with sliding-window phylogenetic analyses, estimating different phylogenetic trees depending on the different regions of the loci. When a locus is important in the color pattern expression, there is a close phylogenetic relationship between the species. When the locus is not important in the color pattern expression, the two species are phylogenetically distant because there is no introgression at such loci.

Domestic species

Introgression can have a significant impact between wild and domestic populations of animals. This includes household pets, as seen in cats [19] or in dogs. [20]

Plants

Introgression has been observed in several plant species. For instance, a species of iris from southern Louisiana has been studied by Arnold and Bennett (1993) regarding the increased fitness of hybrid variants. [21] [22]

Fish

Espinasa et al. found that introgression between a surface-dwelling members of Astroblepus and a troglomorphic species, Astroblepus pholeter, resulted in the development of previously lost traits in offspring, such as distinct eyes and optic nerves. [23]

Introgression line

An introgression line (IL) is a crop species that contains genetic material artificially derived from a wild relative population through repeated backcrossing. An example of a collection of ILs (called an IL-Library) is the use of chromosome segments from Solanum pennellii (a wild species of tomato) that was introgressed into Solanum lycopersicum (the cultivated tomato). The lines of an IL-library usually cover the complete genome of the donor. Introgression lines allow the study of quantitative trait loci, but also the creation of new varieties by introducing exotic traits. [24]

Lineage fusion

Lineage fusion on a phylogenetic tree; individual introgression events within the hybrid zone (blue) are shown as horizontal lines. Hybrids are produced before the lineages fuse, but rejoin one of the two lineages. Neither species A nor B remains extant in the area of interest. LineageFusion.png
Lineage fusion on a phylogenetic tree; individual introgression events within the hybrid zone (blue) are shown as horizontal lines. Hybrids are produced before the lineages fuse, but rejoin one of the two lineages. Neither species A nor B remains extant in the area of interest.

Lineage fusion is an extreme variant of introgression that results from the merging of two distinct species or populations. This eventually results in a single population that displaces or replaces the parental species in the region. [25] Some lineage fusion occurs soon after two taxa diverge or speciate, especially if there are few reproductive barriers between lineages. [26] It is not strictly necessary for the two lineages to be closely related, but rather have the ability to produce viable offspring.

See also

Related Research Articles

Microevolution is the change in allele frequencies that occurs over time within a population. This change is due to four different processes: mutation, selection, gene flow and genetic drift. This change happens over a relatively short amount of time compared to the changes termed macroevolution.

<span class="mw-page-title-main">Hybrid (biology)</span> Offspring of cross-species reproduction

In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different varieties, 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, but can show hybrid vigor, sometimes growing larger or taller than either parent. The concept of 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.

<span class="mw-page-title-main">Polyploidy</span> Condition where cells of an organism have more than two paired sets of chromosomes

Polyploidy is a condition in which the cells of an organism have more than one pair of (homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one from each of two parents; each set contains the same number of chromosomes, and the chromosomes are joined in pairs of homologous chromosomes. However, some organisms are polyploid. Polyploidy is especially common in plants. Most eukaryotes have diploid somatic cells, but produce haploid gametes by meiosis. A monoploid has only one set of chromosomes, and the term is usually only applied to cells or organisms that are normally diploid. Males of bees and other Hymenoptera, for example, are monoploid. Unlike animals, plants and multicellular algae have life cycles with two alternating multicellular generations. The gametophyte generation is haploid, and produces gametes by mitosis; the sporophyte generation is diploid and produces spores by meiosis.

<span class="mw-page-title-main">Gene flow</span> Transfer of genetic variation from one population to another

In population genetics, gene flow is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent allele frequencies and therefore can be considered a single effective population. It has been shown that it takes only "one migrant per generation" to prevent populations from diverging due to drift. Populations can diverge due to selection even when they are exchanging alleles, if the selection pressure is strong enough. Gene flow is an important mechanism for transferring genetic diversity among populations. Migrants change the distribution of genetic diversity among populations, by modifying allele frequencies. High rates of gene flow can reduce the genetic differentiation between the two groups, increasing homogeneity. For this reason, gene flow has been thought to constrain speciation and prevent range expansion by combining the gene pools of the groups, thus preventing the development of differences in genetic variation that would have led to differentiation and adaptation. In some cases dispersal resulting in gene flow may also result in the addition of novel genetic variants under positive selection to the gene pool of a species or population

<i>Cicer</i> Genus of flowering plants

Cicer is a genus of the legume family, Fabaceae, and the only genus found in tribe Cicereae. It is included within the IRLC, and its native distribution is across the Middle East and Asia. Its best-known and only domesticated member is Cicer arietinum, the chickpea.

Backcrossing is a crossing of a hybrid with one of its parents or an individual genetically similar to its parent, to achieve offspring with a genetic identity closer to that of the parent. It is used in horticulture, animal breeding, and production of gene knockout organisms.

<span class="mw-page-title-main">Conservation genetics</span> Interdisciplinary study of extinction avoidance

Conservation genetics is an interdisciplinary subfield of population genetics that aims to understand the dynamics of genes in a population for the purpose of natural resource management, conservation of genetic diversity, and the prevention of species extinction. Scientists involved in conservation genetics come from a variety of fields including population genetics, research in natural resource management, molecular ecology, molecular biology, evolutionary biology, and systematics. The genetic diversity within species is one of the three fundamental components of biodiversity, so it is an important consideration in the wider field of conservation biology.

<span class="mw-page-title-main">Domestication of vertebrates</span>

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<span class="mw-page-title-main">Hybrid zone</span>

A hybrid zone exists where the ranges of two interbreeding species or diverged intraspecific lineages meet and cross-fertilize. Hybrid zones can form in situ due to the evolution of a new lineage but generally they result from secondary contact of the parental forms after a period of geographic isolation, which allowed their differentiation. Hybrid zones are useful in studying the genetics of speciation as they can provide natural examples of differentiation and (sometimes) gene flow between populations that are at some point between representing a single species and representing multiple species in reproductive isolation.

<span class="mw-page-title-main">Grizzly–polar bear hybrid</span> Cross between grizzly and polar bear

A grizzly–polar bear hybrid is a rare ursid hybrid that has occurred both in captivity and in the wild. In 2006, the occurrence of this hybrid in nature was confirmed by testing the DNA of a unique-looking bear who had been shot near Sachs Harbour, Northwest Territories, on Banks Island in the Canadian Arctic. The number of confirmed hybrids has since risen to eight, all of them descending from the same female polar bear.

<span class="mw-page-title-main">Hybrid speciation</span> Form of speciation involving hybridization between two different species

Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was thought to be particularly difficult to achieve, and thus hybrid species were thought to be very rare. With DNA analysis becoming more accessible in the 1990s, hybrid speciation has been shown to be a somewhat common phenomenon, particularly in plants. In botanical nomenclature, a hybrid species is also called a nothospecies. Hybrid species are by their nature polyphyletic.

<i>Heliconius</i> Genus of brush-footed butterflies

Heliconius comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America as far north as the southern United States. The larvae of these butterflies eat passion flower vines (Passifloraceae). Adults exhibit bright wing color patterns which signal their distastefulness to potential predators.

<span class="mw-page-title-main">Genetic pollution</span> Problematic gene flow into wild populations

Genetic pollution is a term for uncontrolled gene flow into wild populations. It is defined as "the dispersal of contaminated altered genes from genetically engineered organisms to natural organisms, esp. by cross-pollination", but has come to be used in some broader ways. It is related to the population genetics concept of gene flow, and genetic rescue, which is genetic material intentionally introduced to increase the fitness of a population. It is called genetic pollution when it negatively impacts the fitness of a population, such as through outbreeding depression and the introduction of unwanted phenotypes which can lead to extinction.

<span class="mw-page-title-main">Hybrid swarm</span> Population of hybrids beyond first hybrid generation

A hybrid swarm is a population of hybrids that has survived beyond the initial hybrid generation, with interbreeding between hybrid individuals and backcrossing with its parent types. Such population are highly variable, with the genetic and phenotypic characteristics of individuals ranging widely between the two parent types. Hybrid swarms thus blur the boundary between the parent taxa. Precise definitions of which populations can be classified as hybrid swarms vary, with some specifying simply that all members of a population should be hybrids, while others differ in whether all members should have the same or different levels of hybridization.

<span class="mw-page-title-main">Secondary contact</span>

Secondary contact is the process in which two allopatricaly distributed populations of a species are geographically reunited. This contact allows for the potential for the exchange of genes, dependent on how reproductively isolated the two populations have become. There are several primary outcomes of secondary contact: extinction of one species, fusion of the two populations back into one, reinforcement, the formation of a hybrid zone, and the formation of a new species through hybrid speciation.

This glossary of genetics and evolutionary biology is a list of definitions of terms and concepts used in the study of genetics and evolutionary biology, as well as sub-disciplines and related fields, with an emphasis on classical genetics, quantitative genetics, population biology, phylogenetics, speciation, and systematics. Overlapping and related terms can be found in Glossary of cellular and molecular biology, Glossary of ecology, and Glossary of biology.

Hybrid incompatibility is a phenomenon in plants and animals, wherein offspring produced by the mating of two different species or populations have reduced viability and/or are less able to reproduce. Examples of hybrids include mules and ligers from the animal world, and subspecies of the Asian rice crop Oryza sativa from the plant world. Multiple models have been developed to explain this phenomenon. Recent research suggests that the source of this incompatibility is largely genetic, as combinations of genes and alleles prove lethal to the hybrid organism. Incompatibility is not solely influenced by genetics, however, and can be affected by environmental factors such as temperature. The genetic underpinnings of hybrid incompatibility may provide insight into factors responsible for evolutionary divergence between species.

Eukaryote hybrid genomes result from interspecific hybridization, where closely related species mate and produce offspring with admixed genomes. The advent of large-scale genomic sequencing has shown that hybridization is common, and that it may represent an important source of novel variation. Although most interspecific hybrids are sterile or less fit than their parents, some may survive and reproduce, enabling the transfer of adaptive variants across the species boundary, and even result in the formation of novel evolutionary lineages. There are two main variants of hybrid species genomes: allopolyploid, which have one full chromosome set from each parent species, and homoploid, which are a mosaic of the parent species genomes with no increase in chromosome number.

<span class="mw-page-title-main">Introgressive hybridization in plants</span>

Introgressive hybridization, also known as introgression, is the flow of genetic material between divergent lineages via repeated backcrossing. In plants, this backcrossing occurs when an generation hybrid breeds with one or both of its parental species.

Hybridization, when new offspring arise from crosses between individuals of the same or different species, results in the assemblage of diverse genetic material and can act as a stimulus for evolution. Hybrid species are often more vigorous and genetically differed than their ancestors. There are primarily two different forms of hybridization: natural hybridization in an uncontrolled environment, whereas artificial hybridization occurs primarily for the agricultural purposes.

References

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Further reading