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The homeodomain is encoded by a 180-base-pair DNA sequence known as the homeobox . The homeodomains appear to be the sites of these proteins that bind DNA, and they are critical in specifying cell fates. For instance, if a chimeric protein is constructed mostly of Antennapedia but with the carboxyl terminus (including the homeodomain) of Ultrabithorax, it can substitute for Ultrabithorax and specify the appropriate cells as parasegment 6 ( Buy Cheap Explore Wholesale Quality Le 3/4 Henley Tee in Blue Frame Denim Release Dates Authentic Prices Cheap Online 4VYEfdkaZ
). The isolated homeodomain of Antennapedia will bind to the same promoters as the entire Antennapedia protein, indicating that the binding of this protein is dependent on its homeodomain ( Prada Ruffled Short Sleeve Top Big Sale 5f7MV
).

The homeodomain folds into three α helices, the latter two folding into a helix-turn-helix conformation that is characteristic of transcription factors that bind DNA in the major groove of the double helix ( Otting et al. 1990 ; Percival-Smith et al. 1990 ). The third helix is the recognition helix, and it is here that the amino acids make contact with the bases of the DNA. A four-base motif, TAAT, is conserved in nearly all sites recognized by homeodomains; it probably distinguishes those sites to which homeodomain proteins can bind. The 5´ terminal T appears to be critical in this recognition, as mutating it destroys all homeodomain binding. The base pairs following the TAAT motif are important in distinguishing between similar recognition sites. For instance, the next base pair is recognized by amino acid 9 of the recognition helix. Mutation studies have shown that the Bicoid and Antennapedia homeodomain proteins use lysine and glutamine, respectively, at position 9 to distinguish related recognition sites. The lysine of the Bicoid homeodomain recognizes the G of CG pairs, while the glutamine of the Antennapedia homeodomain recognizes the A of AT pairs ( Figure 9.32 ; Rachel Zoe Short Sleeve Mini Dress w/ Tags Footlocker Pictures Cheap Online 1pWad
). If the lysine in Bicoid is replaced by glutamine, the resulting protein will recognize Antennapedia-binding sites ( Iro Jalie LeatherTrimmed Skirt w/ Tags Cheap Nicekicks Cheap Sale Release Dates Ts6rE9wLXy
, 1991 ). Other homeodomain proteins show a similar pattern, in which one portion of the homeodomain recognizes the common TAAT sequence, while another portion recognizes a specific structure adjacent to it.

Cofactors for the Hom-C Genes

The genes of the Drosophila homeotic complex specify segmental fates, but they may need some help in doing it. The DNA-binding sites recognized by the homeodomains of the Hom-C proteins are very similar, and there is some overlap in their binding specificity. In 1990, Peifer and Wieschaus discovered that the product of the Extradenticle ( Exd ) gene interacts with several Hom-C proteins and may help specify the segmental identities. For instance, the Ubx protein is responsible for specifying the identity of the first abdominal segment (A1). Without Extradenticle protein, it will transform this segment into A3. Moreover, the Exd and Ubx proteins are both needed for the regulation of the decapentaplegic gene, and the structure of the decapentaplegic promoter suggests that the Extradenticle protein may dimerize with the Ubx protein on the enhancer of this target gene ( Sale Nicekicks Nili Lotan MidRise LaceUp Jeans Free Shipping Explore Cheap Sale Explore A75eu
; van Dyke and Murre 1994 ). The Extradenticle protein includes a homeodomain, and the human protein PBX1, which resembles the Extradenticle protein, may play a similar role as a cofactor for human homeotic genes.

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