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Developmental modules based on a common set of genetic instructions can evolve separately within a species because genetic switches control how the toolkit is used. These switches include:
gene promoters and the *transcription factors that bind to promoters.
enhancers and repressors that can modulate the interactions of transcription factors and promoters.
signaling molecules that activate pathways or gene expression.
signal transduction components that mediate signaling and its effects.
*connect the concepts How transcription factors act as inducers or repressors of gene expression is explained in Key Concept 16.2.
Multiple switches control each gene, creating different expression patterns in different locations. In this way, elements of the genetic toolkit can be involved in multiple developmental processes and still allow individual modules to develop and evolve independently.
Animation 19.4 Modularity
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During evolution, changes in the functions of genetic switches have led to changes in the forms and functions of organisms. To illustrate this, let’s look at the development of wings in Drosophila and other insects. Drosophila species are members of the insect group Diptera, which means “two wings”—that is, they have a single pair of wings, whereas most insects have two pairs of wings (i.e., four wings). The single pair of wings of dipterans develops on the second thoracic segment, where the Hox gene Antennapedia (Antp) is expressed. Antp is also expressed in the third thoracic segment, but in that segment a pair of balancing organs called halteres develops in dipterans. A critical difference between thoracic segments 2 and 3 is that another Hox gene, Ultrabithorax (Ubx), is expressed along with Antp in segment 3 (Figure 19.16). Ubx represses Antp function in dipterans. If Ubx is inactivated by mutation, a second pair of wings forms in thoracic segment 3, as is typical of many other insect groups. Thus some major morphological differences among groups of animals can result from relatively small changes in gene expression.
Q: What would happen if Ubx was genetically engineered to be expressed in all thoracic segments in a butterfly embryo?
If Ubx was expressed in all segments, the expression of the wing-