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SELECTION EXPERIMENTS
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Selection experiments are done for several reasons. Plant and animal breeders select the most desirable individuals as parents in order to improve their stock. Population geneticists select speci c characteristics for study in order to understand the nature of quantitative genetic control. For example, Drosophila were tested in a fteenchoice maze for geotactic response ( g. 18.13).The maze was on its side, so at every intersection, a y had to make a choice between going up or going down. The ies with the highest scores were chosen as parents for the high line (positive geotaxis; favored downward direction), and the ies with the lowest score were chosen as parents for the low line (negative geotaxis; favored upward direction). The same selection was made for each generation. As time progressed, the two lines diverged quite signi cantly. This tells us that there is a large genetic component to the response; the experimenters are successfully amassing more of the downward alleles in the high line and more of the upward alleles in the low line. Several other points emerge from this graph. First, the high and low responses are slightly different, or asymmetrical. The high line responded more quickly, leveled out more quickly, and tended toward the original state more slowly after selection was relaxed. (The relaxation of selection occurred when the parents were a random sample of the adults rather than the extremes for geotactic scores.) The low line responded more slowly and erratically. In addition, the low line returned toward the original state more quickly when selection was relaxed. The nature of these responses ( g. 18.13) indicates that the high line became more homozygous than the low line. This is shown by the former s response when selection is relaxed: It has exhausted a good deal of its variability for the polygenes responsible for geotaxis. The low line,
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Tamarin: Principles of Genetics, Seventh Edition
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IV. Quantitative and Evolutionary Genetics
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18. Quantitative Inheritance
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Quantitative Inheritance
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Selection for geotaxis. The dotted lines represent relaxed selection.
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Dobzhansky and B. Spassky, Arti cial and natural selection for two behavioral traits in Drosophila pseudoobscura, Proceedings of the National Academy of Sciences, USA, 62:75 80, 1969.)
however, seems to have much of its original genetic variability, because the relaxation of selection caused the mean score of this line to increase rapidly. It still had enough genetic variability to head back to the original population mean. The response to a selection experiment is one way that plant and animal breeders can predict future response.
breeding.) Thus, breeders need some index of the potential response to selection so that they can then get the greatest amount of selection with the lowest risk of inbreeding depression.
Realized Heritability
Breeders often calculate a heritability estimate, a value that predicts to what extent their selection will be successful. Heritability is de ned in the following equation: H in which H YO Y YP heritability offspring yield mean yield of the population parental yield YO YP Y Y gain selection differential
(18.8)
H E R I TA B I L I T Y
Plant and animal breeders want to improve the yields of their crops to the greatest degree they can. They must choose the parents of the next generation on the basis of this generation s yields; thus, they are continually performing selection experiments. Breeders run into two economic problems. They cannot pick only the very best to be the next generation s parents because (1) they cannot afford to decrease the size of a crop by using only a very few select parents and (2) they must avoid inbreeding depression, which occurs when plants are self-fertilized or animals are bred with close relatives for many generations. After frequent inbreeding, too much homozygosity occurs, and many genes that are slightly or partially deleterious begin to show themselves, depressing vigor and yield. ( 19 presents more on in-
From this equation, we can see that heritability is the gain in yield divided by the amount of selection practiced ( g. 18.14). YO Y is the improvement over the population average due to YP Y , which is the amount of difference between the parents and the population average. If there is no gain ( YO Y ), then the heritability
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