SHATTERPROOF2 is an Arabidopsis thaliana gene encoding a MADS-box transcription factor that acts with SHATTERPROOF1 in fruit dehiscence. The 2000 study connected their combined activity to dehiscence zone differentiation and lignin deposition in adjacent cells.[ref]
SHATTERPROOF MADS-box genes control seed dispersal in Arabidopsis by Sarah J. Liljegren and others. Abstract; mutant and constitutive-expression studies
Loss-of-function and constitutive-expression experiments ask complementary questions: what fails when the normal regulators are absent, and what changes when their activity is supplied outside its usual pattern? Neither question identifies the transcription factor's direct DNA targets by itself. Establishing regulation of tissue development is a different evidential step from identifying each directly bound gene.
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