Outbreeding depression

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In biology, outbreeding depression happens when crosses between two genetically distant groups or populations result in a reduction of fitness. [1] The concept is in contrast to inbreeding depression, although the two effects can occur simultaneously. [2] Outbreeding depression is a risk that sometimes limits the potential for genetic rescue or augmentations. [1] It is considered postzygotic response because outbreeding depression is noted usually in the performance of the progeny. [3]

Contents

Outbreeding depression manifests in two ways:

Mechanisms for generating outbreeding depression

The different mechanisms of outbreeding depression can operate at the same time. However, determining which mechanism is likely to occur in a particular population can be very difficult.

There are three main mechanisms for generating outbreeding depression:

  1. Fixed chromosomal differences resulting in the partial or complete sterility of F1 hybrids. [1]
  2. Adaptive differentiation among populations
  3. Population bottlenecks and genetic drift

Some mechanisms may not appear until two or more generations later (F2 or greater), [5] when recombination has undermined vitality of positive epistasis. Hybrid vigor in the first generation can, in some circumstances, be strong enough to mask the effects of outbreeding depression. An example of this is that plant breeders will make F1 hybrids from purebred strains, which will improve the uniformity and vigor of the offspring, however the F2 generation are not used for further breeding because of unpredictable phenotypes in their offspring. Unless there is strong selective pressure, outbreeding depression can increase in further generations as coadapted gene complexes are broken apart without the forging of new coadapted gene complexes to take their place. If the outcrossing is limited and populations are large enough, selective pressure acting on each generation can restore fitness. Unless the F1 hybrid generation is sterile or very low fitness, selection will act in each generation using the increased diversity to adapt to the environment. [6] This can lead to recovery in fitness to baseline, and sometimes even greater fitness than original parental types in that environment. [7] However, as the hybrid population will likely to go through a decline in fitness for a few generations, they will need to persist long enough to allow selection to act before they can rebound. [8]

Examples

The first mechanism has the greatest effects on fitness for polyploids, an intermediate effect on translocations, and a modest effect on centric fusions and inversions. [1] Generally this mechanism will be more prevalent in the first generation (F1) after the initial outcrossing when most individuals are made up of the intermediate phenotype.

Examples of the second mechanism include stickleback fish, which developed benthic and limnetic forms when separated. When crosses occurred between the two forms, there were low spawning rates. However, when the same forms mated with each other and no crossing occurred between lakes, the spawning rates were normal. This pattern has also been studied in Drosophila and leaf beetles, where the F1 progeny and later progeny resulted in intermediate fitness between the two parents. This circumstance is more likely to happen and occurs more quickly with selection than genetic drift. [1]

For the third mechanism, examples include poison dart frogs, anole lizards, and cichlid fish. Selection over genetic drift seems to be the dominant mechanism for outbreeding depression. [1]

In plants

For plants, outbreeding depression represents a partial crossing barrier. [3] Unfortunately, outbreeding depression is not understood well in angiosperms. After observing Ipomopsis aggregata over time by crossing plants that were between 10–100m apart, a pattern was noticed that plants that were farther away spatially had a higher likelihood of outbreeding depression. [3] Some general takeaways from this were that spatial patterns of selection on plant genotypes will vary in scale and pattern, and outbreeding depression reflects the genetic constitution of "hybrid" progeny and the environments in which the parents and progeny grow. [3] This means that although outbreeding depression cannot be predicted in angiosperms yet, the environment has a role in it.

See also

Related Research Articles

<span class="mw-page-title-main">Inbreeding</span> Reproduction by closely related organisms

Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and other consequences that may arise from expression of deleterious or recessive traits resulting from incestuous sexual relationships and consanguinity. Animals avoid incest only rarely.

Small populations can behave differently from larger populations. They are often the result of population bottlenecks from larger populations, leading to loss of heterozygosity and reduced genetic diversity and loss or fixation of alleles and shifts in allele frequencies. A small population is then more susceptible to demographic and genetic stochastic events, which can impact the long-term survival of the population. Therefore, small populations are often considered at risk of endangerment or extinction, and are often of conservation concern.

<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

<span class="mw-page-title-main">Self-pollination</span> Form of pollination

Self-pollination is a form of pollination in which pollen from the same plant arrives at the stigma of a flower or at the ovule. There are two types of self-pollination: in autogamy, pollen is transferred to the stigma of the same flower; in geitonogamy, pollen is transferred from the anther of one flower to the stigma of another flower on the same flowering plant, or from microsporangium to ovule within a single (monoecious) gymnosperm. Some plants have mechanisms that ensure autogamy, such as flowers that do not open (cleistogamy), or stamens that move to come into contact with the stigma. The term selfing that is often used as a synonym, is not limited to self-pollination, but also applies to other types of self-fertilization.

Heterosis, hybrid vigor, or outbreeding enhancement is the improved or increased function of any biological quality in a hybrid offspring. An offspring is heterotic if its traits are enhanced as a result of mixing the genetic contributions of its parents. The heterotic offspring often has traits that are more than the simple addition of the parents' traits, and can be explained by Mendelian or non-Mendelian inheritance. Typical heterotic/hybrid traits of interest in agriculture are higher yield, quicker maturity, stability, drought tolerance etc.

A heterozygote advantage describes the case in which the heterozygous genotype has a higher relative fitness than either the homozygous dominant or homozygous recessive genotype. Loci exhibiting heterozygote advantage are a small minority of loci. The specific case of heterozygote advantage due to a single locus is known as overdominance. Overdominance is a rare condition in genetics where the phenotype of the heterozygote lies outside of the phenotypical range of both homozygote parents, and heterozygous individuals have a higher fitness than homozygous individuals.

An F1 hybrid (also known as filial 1 hybrid) is the first filial generation of offspring of distinctly different parental types. F1 hybrids are used in genetics, and in selective breeding, where the term F1 crossbreed may be used. The term is sometimes written with a subscript, as F1 hybrid. Subsequent generations are called F2, F3, etc.

<span class="mw-page-title-main">Minimum viable population</span> Smallest size a biological population can exist without facing extinction

Minimum viable population (MVP) is a lower bound on the population of a species, such that it can survive in the wild. This term is commonly used in the fields of biology, ecology, and conservation biology. MVP refers to the smallest possible size at which a biological population can exist without facing extinction from natural disasters or demographic, environmental, or genetic stochasticity. The term "population" is defined as a group of interbreeding individuals in similar geographic area that undergo negligible gene flow with other groups of the species. Typically, MVP is used to refer to a wild population, but can also be used for ex-situ conservation.

<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 populations principally to avoid extinction. Therefore, it applies genetic methods to the conservation and restoration of biodiversity. Researchers involved in conservation genetics come from a variety of fields including population genetics, molecular ecology, biology, evolutionary biology, and systematics. Genetic diversity is one of the three fundamental levels of biodiversity, so it is directly important in conservation. Genetic variability influences both the health and long-term survival of populations because decreased genetic diversity has been associated with reduced fitness, such as high juvenile mortality, diminished population growth, reduced immunity, and ultimately, higher extinction risk.

Genetic load is the difference between the fitness of an average genotype in a population and the fitness of some reference genotype, which may be either the best present in a population, or may be the theoretically optimal genotype. The average individual taken from a population with a low genetic load will generally, when grown in the same conditions, have more surviving offspring than the average individual from a population with a high genetic load. Genetic load can also be seen as reduced fitness at the population level compared to what the population would have if all individuals had the reference high-fitness genotype. High genetic load may put a population in danger of extinction.

<span class="mw-page-title-main">Molecular ecology</span> Field of evolutionary biology

Molecular ecology is a field of evolutionary biology that is concerned with applying molecular population genetics, molecular phylogenetics, and more recently genomics to traditional ecological questions. It is virtually synonymous with the field of "Ecological Genetics" as pioneered by Theodosius Dobzhansky, E. B. Ford, Godfrey M. Hewitt, and others. These fields are united in their attempt to study genetic-based questions "out in the field" as opposed to the laboratory. Molecular ecology is related to the field of conservation genetics.

Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding. Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. Inbreeding depression is often the result of a population bottleneck. In general, the higher the genetic variation or gene pool within a breeding population, the less likely it is to suffer from inbreeding depression, though inbreeding and outbreeding depression can simultaneously occur.

<span class="mw-page-title-main">Captive breeding</span> Of wild organisms, by humans

Captive breeding, also known as captive propagation, is the process of plants or animals in controlled environments, such as wildlife reserves, zoos, botanic gardens, and other conservation facilities. It is sometimes employed to help species that are being threatened by the effects of human activities such as climate change, habitat loss, fragmentation, over hunting or fishing, pollution, predation, disease, and parasitism.

Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derives from the Greek roots philo, "liking, loving" and patra, "fatherland", although in recent years the term has been applied to more than just the animal's birthplace. Recent usage refers to animals returning to the same area to breed despite not being born there, and migratory species that demonstrate site fidelity: reusing stopovers, staging points, and wintering grounds.

Allogamy or cross-fertilization is the fertilization of an ovum from one individual with the spermatozoa of another. By contrast, autogamy is the term used for self-fertilization. In humans, the fertilization event is an instance of allogamy. Self-fertilization occurs in hermaphroditic organisms where the two gametes fused in fertilization come from the same individual. This is common in plants and certain protozoans.

Extinction vortices are a class of models through which conservation biologists, geneticists and ecologists can understand the dynamics of and categorize extinctions in the context of their causes. This model shows the events that ultimately lead small populations to become increasingly vulnerable as they spiral toward extinction. Developed by M. E. Gilpin and M. E. Soulé in 1986, there are currently four classes of extinction vortices. The first two deal with environmental factors that have an effect on the ecosystem or community level, such as disturbance, pollution, habitat loss etc. Whereas the second two deal with genetic factors such as inbreeding depression and outbreeding depression, genetic drift etc.

Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity.

Genetic purging is the reduction of the frequency of a deleterious allele, caused by an increased efficiency of natural selection prompted by inbreeding.

Autogamy, or self-fertilization, refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species of protists have also been observed using autogamy as a means of reproduction. Flowering plants engage in autogamy regularly, while the protists that engage in autogamy only do so in stressful environments.

Reproductive assurance occurs as plants have mechanisms to assure full seed set through selfing when outcross pollen is limiting. It is assumed that self-pollination is beneficial, in spite of potential fitness costs, when there is insufficient pollinator services or outcross pollen from other individuals to accomplish full seed set.. This phenomenon has been observed since the 19th century, when Darwin observed that self-pollination was common in some plants. Constant pollen limitation may cause the evolution of automatic selfing, also known as autogamy. This occurs in plants such as weeds, and is a form of reproductive assurance. As plants pursue reproductive assurance through self-fertilization, there is an increase in homozygosity, and inbreeding depression, due to genetic load, which results in reduced fitness of selfed offspring. Solely outcrossing plants may not be successful colonizers of new regions due to lack of other plants to outcross with, so colonizing species are expected to have mechanisms of reproductive assurance - an idea first proposed by Herbert Baker and referred to as Baker's Law. Baker’s Law predicts that reproductive assurance should be common in weedy plants that persist by colonizing new sites. As plants evolve towards increase self-fertilization, energy is redirected to seed production rather than characteristics that increased outcrossing, such as floral attractants, which is a condition known as the selfing syndrome.

References

PD-icon.svg This article incorporates public domain material from Michael Lynch. Inbreeding depression and outbreeding depression. National Park Service.
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