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Flower development of Lilium longiflorum - The Lilium information ...

Flower development of Lilium longiflorum - The Lilium information ...

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Floral homeotic mutants <strong>of</strong> <strong>Lilium</strong><br />

Regulatory elements <strong>of</strong> transcription and the ABCDE model for flower<br />

<strong>development</strong>.<br />

Transcription <strong>of</strong> <strong>development</strong>al genes are under the regulation <strong>of</strong> trans and cis-<br />

regulatory elements that dictate the exact moment the genes are turned on and <strong>of</strong>f in<br />

order to trigger a proper and regular <strong>development</strong>al pattern for each species.<br />

Trans-acting elements are protein factors operating in gene promoter regions<br />

(i.e. cis-regulatory regions) in order to be associated with other factors in the RNA<br />

polymerase complex and activate transcription or, alternatively, to inhibit the<br />

transcription <strong>of</strong> a given gene. Transcription factors, such as the MADS proteins, act as<br />

trans-acting elements in eukaryotic organisms.<br />

Cis-regulatory regions are boxes present in gene promoters and other<br />

regulatory regions that modulate the spatial and temporal transcription pattern <strong>of</strong> a<br />

given gene by allowing selective interaction with transcription factors (i.e. trans-acting<br />

elements) and RNA polymerase. <strong>The</strong> CArG box is a known cis-regulatory element<br />

present in promoters <strong>of</strong> genes that interact with the MADS domain in order to assist<br />

their transcription (Pollock and Treisman, 1991).<br />

<strong>The</strong> ABCDE model is mostly composed <strong>of</strong> MADS-box transcription factors<br />

ruling as key elements to activate (or repress) specific genes in flower <strong>development</strong><br />

that will ultimately result in formation <strong>of</strong> floral organs. Changes in the sequence <strong>of</strong><br />

their cis-regulatory regions or trans-acting elements may result in genetic misfunction,<br />

leading to homeotic changes in the flower pattern.<br />

Lily flowers with their homeotic identity partially lost in the third whorl, as<br />

shown in Figure 1b, can be easily found. This change can be triggered by<br />

environmental conditions, suggesting a form <strong>of</strong> epigenetic regulation <strong>of</strong> homeotic<br />

genes involved in flower organ <strong>development</strong>. In the festiva phenotype, however, the<br />

homeotic change <strong>of</strong> the third whorl organs is regular and complete, suggesting a more<br />

severe cause than those triggered by stress conditions.<br />

In this way, if a specific element regulating the expression <strong>of</strong> a C functional<br />

gene is affected, in cis or in trans, its transcription may be suppressed specifically in<br />

the third whorl and not in the fourth whorl, which would lead to specific homeotic<br />

changes <strong>of</strong> stamens into petals (or tepals, in the case <strong>of</strong> Liliaceae). Keeping in mind<br />

that A and C genes have antagonistic functions, alternatively, an extension <strong>of</strong> A-type<br />

gene domain that allows its transcription in the third whorl would repress the C<br />

function and raise specific homeotic changes <strong>of</strong> stamens in petals.<br />

Important cis-regulatory regions <strong>of</strong> AG that control its spatial expression<br />

pattern were found in the second large intron <strong>of</strong> the gene (Sieburth and Meyerowitz,<br />

86

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