Key Points
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The success of Drosophila melanogaster as a model organism is largely due to the power of forward genetic screens to identify the genes that are involved in a biological process. Although traditional genetic screens, such as zygotic lethal screens, have been useful for identifying the genes that act early in fly development, more sophisticated and versatile screens have been developed.
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Modifier screens, in which genes are identified by their ability to alter the phenotype of flies that are genetically sensitized for the process of interest, are useful for finding the components of signal-transduction pathways.
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Clonal screens, in which cells that are homozygous for a mutation of interest can be made in an otherwise heterozygous animal through targeted mitotic recombination (using the Flp/FRT system), allow researchers to identify genes that act in a specific tissue at any stage of development. A modification of this technique, in which clones are made in the germ line, helps to find genes that are maternally supplied to the embryo.
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Mutant screens do not always identify all of the genes for which loss-of-function mutations give the phenotype of interest, as with genes that have redundant functions. In these cases, the GAL4–UAS system can be used to screen for mis- or overexpression phenotypes.
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The success of a genetic screen depends in large part on its design. As mapping and identifying genes that result from a screen can be very laborious, it is often preferable to sacrifice speed and the number of genes recovered in favour of a phenotype that is directly related to the process of interest.
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Genetic screens will continue to be useful in years to come, even once all genes have been identified. The functional characterization of genes relies on having allelic variants of each gene, which, for the time being, only genetic screens can provide.
Abstract
The success of Drosophila melanogaster as a model organism is largely due to the power of forward genetic screens to identify the genes that are involved in a biological process. Traditional screens, such as the Nobel-prize-winning screen for embryonic-patterning mutants, can only identify the earliest phenotype of a mutation. This review describes the ingenious approaches that have been devised to circumvent this problem: modifier screens, for example, have been invaluable for elucidating signal-transduction pathways, whereas clonal screens now make it possible to screen for almost any phenotype in any cell at any stage of development.
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Acknowledgements
I thank N. Brown, B. Dickson, M. Freeman, E. Hafen, B. Sanson and T. Xu for providing pictures, and S. Bray for her helpful comments on the manuscript. D.StJ. is supported by a Wellcome Trust Principal Fellowship.
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DATABASES
tuberous sclerosis complex 1 and 2
FURTHER INFORMATION
Berkeley Drosophila Genome Project
Nature Reviews Genetics Focus on 'The art and design of genetics screens'
Glossary
- POLYTENE CHROMOSOME
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A giant chromosome that is formed by many rounds of replication of the DNA. The replicated DNA molecules tightly align side-by-side in parallel register, which creates a non-mitotic chromosome that is visible by light microscopy.
- PROTOSTOME–DEUTEROSTOME
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The two principal divisions of animal phyla, based on how the mouth forms in the embryo.
- BALANCER CHROMOSOME
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A chromosome with one or more inverted segments that suppress recombination. They are used as genetic tools because they allow lethal mutations to be maintained without selection.
- MATERNAL-EFFECT MUTATION
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Homozygous-viable mutation that causes little or no phenotype in the mutant mothers, but leads to the development of abnormal offspring.
- IPSILATERAL AXON
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An axon that does not cross the midline.
- SEGMENT-POLARITY GENE
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A gene that is required for anteroposterior patterning within each segment, such as wingless, engrailed and hedgehog.
- OMMATIDIA
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The compound eye of Drosophila is formed from 800 ommatidia, each of which contains eight photoreceptor cells, surrounded by four cone cells that secrete the lens, and seven pigment cells.
- IMAGINAL DISC
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Sac-like infolding of the epithelium in the larva. They give rise to most of the external structures of the adult. Imaginal disc cells are set aside in the embryo and continue to divide until pupation, when they differentiate.
- NOTUM
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The dorsal or upper surface of the thoracic segment of any insect.
- FOLLICLE STEM CELL
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Each ovariole (chambers in the ovary through which the egg passes during development) contains 2–3 follicle stem cells, which produce the somatic follicle cells that surround the chambers. The follicle cells then differentiate into several cell types, including the border cells, which migrate from the anterior of the egg chamber towards the oocyte, where they contribute to the formation of the micropyle. At the end of oogenesis, the follicle cells secrete the eggshell and undergo apoptosis.
- MINI-WHITE GENE
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A truncated version of the white gene that is commonly used as the selectable marker in transformation constructs. One copy of the transgene usually produces yellow or orange eyes in a white mutant background, whereas two copies give more complete rescue and produce darker eye colours. This allows more than one transgene to be followed at a time, and flies that are either heterozygous or homozygous for a particular insertion to be distinguished.
- ISOGENESIS
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A way of homogenizing the genetic background of a line that is used for mutagenesis. In an isogenic stock, the two homologous chromosomes of each pair are identical, which ensures that no recessive lethal allele is present.
- RNA INTERFERENCE
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(RNAi). A process by which double-stranded RNA specifically silences the expression of homologous genes through degradation of their cognate mRNA.
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St Johnston, D. The art and design of genetic screens: Drosophila melanogaster. Nat Rev Genet 3, 176–188 (2002). https://doi.org/10.1038/nrg751
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DOI: https://doi.org/10.1038/nrg751
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