Abstract
The Arabidopsis stoma is a specialized epidermal valve made up of a pair of guard cells around a pore whose aperture controls gas exchange between the shoot and atmosphere. Guard cells (GCs) are produced by a symmetric division of guard mother cells (GMCs). The R2R3-MYB transcription factor FOUR LIPS (FLP) and its paralogue MYB88 restrict the division of a GMC to one. Previously, the upstream regions of several core cell cycle genes were identified as the direct targets of FLP/MYB88, including the B-type cyclin-dependent kinase CDKB1;1 and A2-type cyclin CYCA2;3. Here we show that CDKA;1 is also an immediate direct target of FLP/MYB88 through the binding to cis-regulatory elements in the CDKA;1 promoter region. CDKA;1 activity is required not only for normal GMC divisions but also for the excessive cell overproliferation in flp myb88 mutant GMCs. The impaired defects of GMC division in cdkb1;1 1;2 mutants could be partially rescued by a stage-specific expression of CDKA;1. Although targeted overexpression of CDKA;1 does not affect stomatal development, ectopic expression of the D3-type cyclin CYCD3;2 induces GC subdivision, resulting in a stoma with 3-4 GCs instead of the normal two. Co-overexpression of CDKA;1 with CYCD3;2, but not with CYCA2;3, confers a synergistic effect with respect to GC subdivision. Thus, in addition to a role in stomatal formative asymmetric divisions at early developmental stages, CDKA;1 is needed in triggering GMC symmetric divisions at the late stage of stomatal development. However, timely down-regulation of CDKA;1-CYCD3 activity is required for restriction of GC proliferation.
Highlights
Guard cell vitality is checked by neutral red staining
Neutral red dyes are taken by living cells and stored in vacuoles
All GCs are vital in stomata from FAMA:amiRNA-RBR1 (D) and FAMA:CYCD3;2 (E)
Summary
Lane1, probes for binding Element A with His-MYB88 proteins; Lane 2, probes for Element B with His-MYB88 proteins; Lane 3, probes for Element C and D Guard cell vitality is checked by neutral red staining.
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