Floral development is one of the model systems for investigating the mechanisms underlying organogenesis in plants. Floral organ identity is controlled by the well-known ABC model, which has been generalized to many flowering plants. Here, we report a previously uncharacterized MYB-like gene, AGAMOUS-LIKE FLOWER (AGLF), involved in flower development in the model legume Medicago truncatula. Loss-of-function of AGLF results in flowers with stamens and carpel transformed into extra whorls of petals and sepals. Compared with the loss-of-function mutant of the class C gene AGAMOUS (MtAG) in M. truncatula, the defects in floral organ identity are similar between aglf and mtag, but the floral indeterminacy is enhanced in the aglf mutant. Knockout of AGLF in the mutants of the class A gene MtAP1 or the class B gene MtPI leads to an addition of a loss-of-C-function phenotype, reflecting a conventional relationship of AGLF with the canonical A and B genes. Furthermore, we demonstrate that AGLF activates MtAG in transcriptional levels in control of floral organ identity. These data shed light on the conserved and diverged molecular mechanisms that control flower development and morphology among plant species.flower development | ABC model | AGAMOUS | Medicago truncatula O rgan differentiation is crucial in the development of living organisms, such as limb-bud growth in mammals, bodysegment development in insects, and flower-organ identity in plants. Most flowers of higher plants consist of four concentric whorls of floral organs: sepals, petals, stamens, and innermost carpel(s). Floral organs have distinct morphologies and functions. The outer sepals and petals provide protection to the reproductive organs inside, i.e., the stamens and carpels that produce gametophytes. The developmental process of four whorls of flower is controlled by the organ identity genes in the well-known ABC model (1-4). In Arabidopsis thaliana, sepals are controlled by the class A genes APETALA 1 (AP1) and AP2; petals are controlled by the class A genes and class B genes AP3 and PISTILLATA (PI); stamens are controlled by the class B gene and class C gene AGAMOUS (AG); and carpels are controlled by the class C gene only (5-9). In another model plant, Antirrhinum majus, similar genes have been identified, such as SQUAMOSA, LIPLESS1, and LIPLESS2 as class A genes, DEFICIENS and GLOBOSA as class B genes, and PLENA and FARINELLI as class C genes (10-12). All of the ABC genes except AP2 and its homologs belong to the MADS (MCM1, AGAMOUS, DEFICIENS A, and SERUM RESPONSE FAC-TOR) homeotic gene family (13). The knockout mutants of the ABC genes display homeotic transformation of the floral organs in corresponding whorls in Arabidopsis. The mutation of AP1 results in the transformation of sepals into carpels and petals into stamens, while the pi mutant causes petals and stamens to be transformed into sepals and carpels. The ag mutant develops indeterminate flowers with stamens and carpel transformed into numerous whorls of petals and sepals.The ABC model ha...