Complementation (genetics) - Wikipedia, the free encyclopedia

http://en.wikipedia.org/wiki/Complementation_(genetics)

Complementation (genetics)
From Wikipedia, the free encyclopedia

In genetics, complementation refers to a relationship between two genetically distinct strains of an organism which both have homozygous recessive mutations that produce the same phenotype (for example, a change in wing structure in flies) but which are not the result of the same allele(s) of a specific gene. If, when these strains are crossed with each other, some offspring show recovery (often termed "rescue") of the mutant phenotype, and thus a return to the wild-type phenotype, then these strains are said to show "genetic complementation". When this occurs, each strain's haploid genome supplies a single wild-type allele to "complement" the mutated allele of the other strain's haploid genome, causing the offspring to have heterozygous genotypes in all related genes. Since the mutations are recessive, the offspring will display the wild-type phenotype. A complementation test (sometimes called a "cis-trans" test) can be used to test whether two strains of a particular gene are complements of one another and was developed by American geneticist Edward B. Lewis. It answers the question: "Does a wild-type copy of gene X rescue the function of the mutant allele that is believed to define gene X?". If there is an allele with an observable phenotype whose function can be provided by a wild type genotype (i.e., the allele is recessive), one can ask whether the function that was lost because of the recessive allele can be provided by another mutant genotype. If not, the two alleles must be defective in the same gene. The convenience and essence of this test is that the trait can serve to describe the gene's function without the exact knowledge of what the gene is doing on a molecular level.[1] Complementation arises because loss of function in genes responsible for different steps in the same metabolic pathway can give rise to the same phenotype. When strains are bred together, offspring inherit wildtype versions of each gene from either parent. Because the mutations are recessive, there is a recovery of function in that pathway, so offspring recover the wild-type phenotype. Thus, the test is used to decide if two independently derived recessive mutant phenotypes are caused by mutations in the same gene or in two different genes. If both parent strains have mutations in the same gene, no normal versions of the gene are inherited by offspring; they express the same mutant phenotype and complementation has failed to occur. In other words: If the combination of two haploid genomes containing different recessive mutations yields a mutant phenotype, then there are three possibilities: 1. Mutations occur in the same gene. 2. One mutation affects the expression of the other. 3. One mutation may result in an inhibitory product. If the combination of two haploid genomes containing different recessive mutations yields the wild type phenotype, then the mutations must be in different genes.

Contents
1 Example of a Simple Complementation Test 2 Complementation tests in fungi and bacteriophage 3 Genetic complementation, heterosis and the evolution of sexual reproduction 4 Exceptions 5 See also

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Complementation (genetics) - Wikipedia, the free encyclopedia

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6 References

Example of a Simple Complementation Test

For a simple example of a complementation test, suppose a geneticist is interested in studying two strains of white-eyed flies of the species Drosophila melanogaster. In this species, wild type flies have red eyes and eye color is known to be related to two genes, A and B. Each one of these genes has two alleles, a dominant one that codes for a working protein (A and B respectively) and a recessive one that codes for a malfunctioning protein (a and b respectively). Since both proteins are necessary for the synthesis of red pigmentation in the eyes, if a given fly is homozygous for either a or b, it will have white eyes. Knowing this, the geneticist may perform a complementation test on two separately obtained strains of pure-breeding white-eyed flies. The test is performed by crossing two flies, one from each strain. If the resulting progeny have red eyes, the two strains are said to complement; if the progeny have white eyes, they do not. If the strains complement, we imagine that one strain must have a genotype aa BB and the other AA bb, which when crossed yield the genotype AaBb. In other words, each strain is homozygous for a different deficiency that produces the same phenotype. If the strains do not complement, they both must have genotypes aa BB, AA bb, or aa bb. In other words, they are both homozygous for the same deficiency, which obviously will produce the same phenotype.
Example of a complementation test. Two strains of flies are white eyed because of two different autosomal recessive mutations which interrupt different steps in a single pigment-producing metabolic pathway. Flies from Strain 1 have complementary mutations to flies from Strain 2 because when they are crossed the offspring are able to complete the full metabolic pathway and thus have red eyes.

Complementation tests in fungi and bacteriophage

Complementation tests can also be carried out with haploid eukaryotes such as fungi, with bacteria and with viruses such as bacteriophage.[2] Research on the fungus Neurospora crassa led to the development of the one-gene-one enzyme concept that provided the foundation for the subsequent development of molecular genetics.[3][4] The complementation test was one of the main tools used in the early Neurospora work, because it was easy to do, and allowed the investigator to determine whether any two nutritional mutants were defective in the same, or different genes. The complementation test was also used in the early development of molecular genetics when bacteriophage T4 was one of the main objects of study.[5] In this case the test depends on mixed infections of host bacterial cells with two different bacteriophage mutant types. Its use was key to defining most of the genes of the virus, and provided the foundation for the study of such fundamental processes as DNA replication and repair, and how molecular machines are constructed.
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Complementation (genetics) - Wikipedia, the free encyclopedia

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Genetic complementation, heterosis and the evolution of sexual reproduction

Heterosis is the tendency for hybrid individuals to exceed their pure bred parents in size and vigor. The phenomenon has long been known in animals and plants. Heterosis appears to be largely due to genetic complementation, that is the masking of deleterious recessive alleles in hybrid individuals. In general, the two fundamental aspects of sexual reproduction in eukaryotes are meiosis, and outcrossing. These two aspects have been proposed to have two natural selective advantages, respectively. Meiosis is proposed to be adaptive because it facilitates recombintional repair of DNA damages that are otherwise difficult to repair (seeMeiosis#Theory that DNA repair is the adaptive advantage of meiosis). Outcrossing is proposed to be adaptive because it facilitates complementation, that is the masking of deleterious recessive alleles [6] (also see Heterosis#Genetic and epigenetic bases of heterosis). The benefit of masking deleterious alleles 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. Further, the selective advantage of complementation that arises from outcrossing may largely account for the general avoidance of inbreeding in nature (e.g see articles Kin recognition, Inbreeding depression and Incest taboo).

Exceptions
There are exceptions to these rules. Two non-allelic mutants may occasionally fail to complement (called "non-allelic non-complementation" or "unlinked non-complementation"). This situation is rare and is dependent on the particular nature of the mutants being tested. For example, two mutations may be synthetically dominant negative. Another exception is transvection, in which the heterozygous combination of two alleles with mutations in different parts of the gene complement each other to rescue a wild type phenotype.

See also
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References
1. ^ Genetics lectures 1-3 '03 (http://ocw.mit.edu/NR/rdonlyres/Biology/7-03Fall-2004/233B3544-E379-44C5BC31-EF7D864AE73B/0/lecture2.pdf) 2. ^ Fincham JRS (1966). Genetic Complementation (http://books.google.com.au/books?id=hdc9AAAAIAAJ) . Microbial and molecular biology. 3. W.A. Benjamin. pp. 118. ASIN B009SQ0G9C (//www.amazon.com /dp/B009SQ0G9C) . OCLC 239023 (//www.worldcat.org/oclc/239023) . http://books.google.com.au /books?id=hdc9AAAAIAAJ. 3. ^ Beadle GW (2007). "Biochemical genetics: Some recollections" (http://books.google.com.au /books?id=g_JSw4LVKtIC&lpg=PR3&dq=ISBN%3A0879698004&pg=PA23) . In Cairns, J.; Stent, G.S.; Watson, J.D.. Phage and the Origins of Molecular Biology (4th ed.). Cold Spring Harbor Laboratory of Quantitative Biology. pp. 2332. ISBN 0879698004. http://books.google.com.au/books?id=g_JSw4LVKtIC&lpg=PR3& dq=ISBN%3A0879698004&pg=PA23. 4. ^ Horowitz NH (April 1991). "Fifty years ago: the Neurospora revolution" (http://www.genetics.org /cgi/pmidlookup?view=long&pmid=1827628) . Genetics 127 (4): 6315. PMC 1204391 (//www.ncbi.nlm.nih.gov /pmc/articles/PMC1204391) . PMID 1827628 (//www.ncbi.nlm.nih.gov/pubmed/1827628) . http://www.genetics.org /cgi/pmidlookup?view=long&pmid=1827628. 5. ^ Epstein RH, Bolle A, Steinberg CM, Kellenberger E, Boy De La Tour E, Chevalley R, Edgar RS, Susman M,

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Complementation (genetics) - Wikipedia, the free encyclopedia

http://en.wikipedia.org/wiki/Complementation_(genetics)

Denhardt GH, Lielausis A (1963). "Physiological studies of conditional lethal mutants of bacteriophage T4D". Cold Spring Harbor Symp. Quant. Biol. 28: 375394. doi:10.1101/SQB.1963.028.01.053 (http://dx.doi.org /10.1101%2FSQB.1963.028.01.053) . 6. ^ Bernstein H, Byerly HC, Hopf FA, Michod RE (September 1985). "Genetic damage, mutation, and the evolution of sex" (http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=3898363) . Science 229 (4719): 127781. PMID 3898363 (//www.ncbi.nlm.nih.gov/pubmed/3898363) . http://www.sciencemag.org /cgi/pmidlookup?view=long&pmid=3898363.

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