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In homeotic mutants, one cell type follows the developmental pathway other cell types normally follow. These genes de ne the future development of segments based on the pattern of expression of the segmentation genes before them. When they mutate, they switch the development of that segment to an adjacent segment, usually anterior to it. Homeotic genes are also called memory genes because they set the developmental fate of a segment, a fate that is
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(c) Figure 16.16 Photos a e show how the margins of expression of two gap genes, fushi tarazu (brown) and even-skipped (gray), narrow and sharpen as time goes on (between about hours 3 and 4 of embryonic development). The stripes appear from staining with antibodies against the proteins. (From P. A. Lawrence, The making of a y, Blackwell Publications, 1992.)
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III. Molecular Genetics
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16. Gene Expression: Control in Eukaryotes
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remembered from one cell division to the next. They are also called master-switch genes since they control the activity of many other genes. Two major homeotic gene complexes are known in Drosophila melanogaster ( g. 16.17): the bithorax complex (BX-C), analyzed extensively by E. Lewis, D. Hogness, and their colleagues, and the Antennapedia complex (ANT-C), worked on extensively by W. Gehring,T. Kaufman, and their colleagues. (The two regions together are known as the Hom-C region.) Genes in the Antennapedia complex control the fate of the anterior develop-
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ment of the fruit y (head and anterior thorax), whereas genes in the bithorax complex control the fate of posterior development (posterior thorax and abdomen). Mutations in the genes of these complexes can change the fate of development of whole sections of the y. For example, Nasobemia, an Antennapedia-complex mutant, causes legs to grow where antennae would normally be located ( g. 16.18); and bithorax, a bithorax-complex mutant, produces ies with two thoraxes (four-winged diptera; g. 16.19). The genes in these complexes are arranged in order of their progressive action from anterior to posterior on the y (see g. 16.17). One model of action for these genes suggests that they require the action of the genes of the adjacent anterior segment plus the action of that homeotic gene itself. Thus, loss of function of a particular gene by mutation would cause a segment to develop like the previous section in the anterior direction.
The Homeo Box
Using recombinant DNA techniques, W. Gehring and his colleagues found a consensus sequence of 180 base pairs of DNA in genes of the Antennapedia and bithorax complexes. Further probing localized this same segment of 180 base pairs to about a dozen genes in Drosophila, all with homeotic or segmentation properties. They thus called this DNA sequence the homeo box. The nucleotides of the homeo box are translated into a peptide region of 60 amino acids called the homeo domain ( g. 16.20, box 16.1). Using a recombinant probe for the homeo box, or a computer search for the consensus sequence, researchers found it in the genes of plants, yeast, sea urchins, frogs, and human beings. This high degree of se-
Edward B. Lewis (1918 ). (Courtesy of
Dr. Edward B. Lewis.)
Walter J. Gehring (1939 ). (Courtesy Dr.
Walter J. Gehring.)
Figure 16.17 A map of the homeotic complexes ANT-C and BX-C in Drosophila, and the regions of the body in which the genes are expressed, mapped on a ten-hour embryo. Note that the genes are expressed from right to left, or in an anterior to posterior direction. Dotted lines indicate lack of detectable function at this stage in development. Embryonic segments are intercalary (int), maxillary (max), labial (lab), thoracic (T1 T3), and abdominal (A1 A8). Genes are labial (lab), proboscipedia (pb), Zerkn llt (Zen), Deformed (Dfd), Sex combs reduced (Scr), and Antennapedia (Antp) in ANT-C and Ultrabithorax (Ubx), abdominal-A (abd-A), and Abdominal-B (Abd-B) in BX-C. Note that Zen is unique in specifying information for the dorsalventral axis rather than the anterior-posterior axis. (Reprinted
from Cell, Vol. 68, W. McGinnis and R. Krumlauf, pp. 283 302, Copyright 1992, with permission from Elsevier Science.)
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