Abscission is the process by which plants shed unwanted organs, either as part of a natural developmental program or in response to environmental stimuli. Studies in Arabidopsis thaliana have elucidated a number of the genetic components that regulate abscission of floral organs, including a pair of related receptor-like protein kinases, HAESA and HAESA-like 2 (HAE/HSL2) that regulate a MAP kinase cascade that is required for abscission. HAE is transcriptionally up-regulated in the floral abscission zone just before cell separation. Here, we identify AGAMOUS-like 15 (AGL15; a MADS-domain transcription factor) as a putative regulator of HAE expression. Overexpression of AGL15 results in decreased expression of HAE as well as a delayed abscission phenotype. Chromatin immunoprecipitation experiments indicate that AGL15 binds the HAE promoter in floral receptacles. AGL15 is then differentially phosphorylated through development in floral receptacles in a MITOGEN-ACTIVATED PROTEIN KINASE KINASE 4/5-dependent manner. MAP kinase phosphorylation of AGL15 is necessary for full HAE expression, thus completing a positive feedback loop controlling HAE expression. Together, the network components in this positive feedback loop constitute an emergent property that regulates the large dynamic range of gene expression (27-fold increase in HAE) observed in flowers when the abscission program is initiated. This study helps define the mechanisms and regulatory networks involved in a receptor-mediated signaling pathway that controls floral organ abscission.A bscission is the process that plants use to shed unwanted organs. Various plants can abscise leaves, fruits, and flowers. One of the most noticeable abscission events occurs when deciduous trees and shrubs shed their leaves in the fall. Plants can abscise organs as part of a developmental program or in an inducible manner in response to stimuli like abiotic or biotic stress. For example, tomatoes can abscise leaves and flowers in response to drought or insect feeding (1-3). In order for abscission to occur, a layer of small and cytoplasmically dense cells, known as an abscission zone, must be laid down during development at the boundary between the organ to be abscised and the body of the plant (4).Arabidopsis thaliana has been used to elucidate a number of the molecular and genetic components that regulate abscission of floral organs. In Arabidopsis, floral organs abscise shortly after pollination, which corresponds to stage-16 flowers or approximately floral position 4-6, where anthesis is defined as position 1 and older flowers are defined as increasing numerical positions ( Fig. 1 A and B) (5). A number of genetic components regulating this abscission process have been described thoroughly in recent reviews (6, 7). In brief, a pair of related receptor-like protein kinases, HAESA and HAESA-like 2 (HAE/HSL2), are required for floral abscission (Fig. 1C) and are thought to be triggered by a peptide derived from INFLORESCENCE DEFICIENT IN ABSCISSION (IDA) (8, 9). A mitogen-act...