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In the  Arabidopsis thaliana  seed pod, pod shatter and seed dispersal properties are in part determined by the development of a longitudinally orientated dehiscence zone (DZ) that derives from cells of the gynoecial valve margin (VM). Transcriptional regulation of the MADS protein encoding transcription factors genes  SHATTERPROOF1  ( SHP1 ) and  SHATTERPROOF2  ( SHP2 ) are critical for proper VM identity specification and later on for DZ development. Current models of  SHP1  and  SHP2  regulation indicate that the transcription factors FRUITFULL (FUL) and REPLUMLESS (RPL) repress these  SHP  genes in the developing valve and replum domains, respectively. Thus the expression of the  SHP  genes is restricted to the VM.  FUL  encodes a MADS-box containing transcription factor that is predicted to act through CArG-box containing  cis -regulatory motifs. Here we delimit functional modules within the  SHP2  cis-regulatory region and examine the functional importance of CArG box motifs within these regulatory regions. We have characterized a 2.2kb region upstream of the  SHP2  translation start site that drives early and late medial domain expression in the gynoecium, as well as expression within the VM and DZ. We identified two separable, independent  cis -regulatory modules, a 1kb promoter region and a 700bp enhancer region, that are capable of giving VM and DZ expression. Our results argue for multiple independent  cis -regulatory modules that support  SHP2  expression during VM development and may contribute to the robustness of  SHP2  expression in this tissue. Additionally, three closely positioned CArG box motifs located in the  SHP2  upstream regulatory region were mutated in the context of the 2.2kb reporter construct. Mutating simultaneously all three CArG boxes caused a moderate de-repression of the  SHP2  reporter that was detected within the valve domain, suggesting that these CArG boxes are involved in  SHP2  repression in the valve.

Redundant CArG Box Cis-motif Activity Mediates SHATTERPROOF2 Transcriptional Regulation during Arabidopsis thaliana Gynoecium Development