Flower development of Lilium longiflorum - The Lilium information ...
Flower development of Lilium longiflorum - The Lilium information ...
Flower development of Lilium longiflorum - The Lilium information ...
Create successful ePaper yourself
Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.
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