Outbreeding depression

In biology, outbreeding depression is when progeny, resulting from crosses between genetically distant individuals (outcrossing), exhibit lower fitness in the parental environment than either of their parents, or than progeny from crosses between individuals that are more closely related.^[1] The concept is opposed to inbreeding depression, although the two effects can occur simultaneously.^[2] Outbreeding depression manifests most significantly in two ways:

• Intermediate genotypes are not adapted to either parental habitat. For example, selection in one population might favor a large body size, whereas in another population small body size might be more advantageous, while individuals with intermediate body sizes are comparatively disadvantaged in both populations. As another example, in the Tatra Mountains, the introduction of Ibex from the Middle East, resulted in hybrids which produce calves at the coldest time of the year.^[3]
• Breakdown of biochemical or physiological compatibility. Within isolated breeding populations, alleles are selected in the context of the local genetic background. Because the same alleles may have rather different effects in different genetic backgrounds—there is the potential evolution of different locally adapted gene complexes. Outcrossing between individuals with differently adapted gene complexes can result in disruption of this selective advantage, resulting in a loss of fitness.

Mechanism and Impact

The different mechanisms of outbreeding depression can operate at the same time. However, determining which mechanism is more important in a particular population is very difficult. Generally the first mechanism will be more prevalent in the first generation (F1) after the initial outcrossing when most individuals are made up of the intermediate phenotype. An extreme case of this type of outbreeding depression is the sterility and other fitness-reducing effects often seen in interspecific hybrids (such as mules), which involves not only different alleles of the same gene but even different orthologous genes.

The second mechanism may not appear until two or more generation later (F2 or greater),^[4] when recombination has undermine vitality of positive epistasis and/or increasing negative 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 F1 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 co-adapted gene complexes are broken apart without the forging of new co-adapted 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.^[5] This can lead to recovery in fitness to baseline, and sometimes even greater fitness than original parental types in that environment.^[6] 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 ^[7]

See also

• Outbreeding
• Heterozygote advantage
• Dominance versus overdominance
• Haldane's Rule
• Miscegenation
• Heterosis
• Inbreeding depression

References

 This article incorporates public domain material from the National Park Service document "Inbreeding depression and outbreeding depression" by Michael Lynch.

1. Leimu, R.; Fischer, M. (2010). Bruun, Hans Henrik, ed. "Between-Population Outbreeding Affects Plant Defence". PLoS ONE. 5 (9): e12614. doi:10.1371/journal.pone.0012614. PMC 2935481. PMID 20838662.
2. Frankham, Ballou, & Briscoe, R., J.D. & D.A. (2002). Introduction to Conservation Genetics. Cambridge. p. 382. ISBN 0521702712.
3. Turcek, FJ (1951). "Effect of introductions on two game populations in Czechoslovakia.". Journal of Wildlife Management 15, 113-114.
4. Fenster, Charles (2000). "Inbreeding and Outbreeding Depression in Natural Populations of Chamaecrista fasciculata (Fabaceae)". Conservation Biology. doi:10.1046/j.1523-1739.2000.99234.x.
5. Erickson and Fenster (2006). "Intraspecific hybridization and the recovery of fitness in the native legume Chamaecrista fasciculata.". Evolution. JSTOR 4095211. PMID 16610315.
6. Lewontin & Birch, R.C. & L.C. (February 3, 1966). "Hybridization as a source of variation for adaptation to new environments". Evolution.
7. Frankham, Ballou, & Briscoe, R., J.D. & D.A. (2002). Introduction to Conservation Genetics. Cambridge. p. 388 ISBN 0521702712