Floral organ abscission in Arabidopsis thaliana is regulated by the putative ligand-receptor system comprising the signaling peptide INFLORESCENCE DEFICIENT IN ABSCISSION (IDA) and the two receptor-like kinases HAESA and HAESA-LIKE2. The IDA signaling pathway presumably activates a MITOGEN-ACTIVATED PROTEIN KINASE (MAPK) cascade to induce separation between abscission zone (AZ) cells. Misexpression of IDA effectuates precocious floral abscission and ectopic cell separation in latent AZ cell regions, which suggests that negative regulators are in place to prevent unrestricted and untimely AZ cell separation. Through a screen for mutations that restore floral organ abscission in ida mutants, we identified three new mutant alleles of the KNOTTED-LIKE HOMEOBOX gene BREVIPEDICELLUS (BP)/KNOTTED-LIKE FROM ARABIDOPSIS THALIANA1 (KNAT1). Here, we show that bp mutants, in addition to shedding their floral organs prematurely, have phenotypic commonalities with plants misexpressing IDA, such as enlarged AZ cells. We propose that BP/KNAT1 inhibits floral organ cell separation by restricting AZ cell size and number and put forward a model whereby IDA signaling suppresses BP/KNAT1, which in turn allows KNAT2 and KNAT6 to induce floral organ abscission.
INTRODUCTIONCell separation processes are critical for the development of a plant and play key roles from sculpting the form of the plant to scattering seeds. Abscission, a physiological process that involves programmed changes in cellular adhesion, allows the plant to discard nonfunctional or infected organs. At the cellular level, the presence of an abscission zone (AZ), consisting of small densely cytoplasmic cells at the boundary between organ and plant, is a prerequisite for abscission to take place (McKim et al., 2008). During the subsequent activation of the cell separation process, AZ cells acquire competence to respond to abscission signals and secrete cell wall-modifying and hydrolyzing enzymes that act to degrade the middle lamella between two adjacent cell files (Bleecker and Patterson, 1997;Patterson, 2001;Roberts et al., 2002;Lewis et al., 2006;Stenvik et al., 2006). Shortly before organ shedding, the cells at the proximal side of the AZ expand; however, the relationship between AZ cell enlargement and organ separation is unclear (Patterson, 2001). It has been proposed that the functional role of AZ cell expansion might be to create the tension needed for the final mechanical rupture of the AZ (Sexton and Redshaw, 1981); indeed, the receptor-like kinase (RLK) EVERSHED (EVR) has recently been implicated in regulating the proper timing of floral organ abscission in Arabidopsis thaliana in part by restricting AZ cell size .The correct temporal and spatial regulation of abscission is crucial during plant development. Premature abscission of reproductive organs or immature seeds can compromise reproduction, and unrestricted cell separation can lead to shedding of organs in the proximity of activated AZ cells or interfere with tissue integrity. Studies using Arabidopsis...