The mechanism of floral organ specification is principally conserved in angiosperms, as demonstrated by the ABC model. By contrast, mechanisms that regulate the development of organs or structures specific to a group of species remain unclear. Grasses have unique inflorescence units, comprising spikelets and florets. In the genus Oryza (rice), the single spikelet consists of a fertile floret subtended by a lemma and a palea, two sterile lemmas, and rudimentary glumes. Each sterile lemma is a tiny glume-like organ with a smooth surface. Here, we have examined a long sterile lemma1 (g1) mutant, in which the sterile lemma is enlarged like the lemma. Detailed phenotypic analysis reveals that the large sterile lemma in the g1 mutant appears to be caused by homeotic transformation of the sterile lemma into a lemma, suggesting that G1 is involved in the repression of lemma identity to specify the sterile lemma. Gene isolation reveals that G1 is a member of a plant-specific gene family that encodes proteins with a previously uncharacterized domain, named here ALOG (Arabidopsis LSH1 and Oryza G1). G1 mRNA is expressed in sterile lemma primordia throughout their development, and G1 protein is localized in the nucleus. A trans-activation assay using the yeast GAL4 system suggests that G1 is involved in transcriptional regulation. Repression of lemma identity by G1 is consistent with a hypothesis proposed to explain the morphological evolution of rice spikelets. We also show that a wild rice species, Oryza grandiglumis, that forms large sterile lemmas has serious mutations in the G1 gene.flower ͉ grass ͉ Oryza ͉ ALOG ͉ morphological evolution A lthough angiosperms produce diverse forms of flowers, from beautiful and entomophilous to inconspicuous and anemophilous, the genetic programs directing floral development are fundamentally conserved. Floral organ specification is explained by the ABC model, and genes constituting this model are likely to be functionally conserved in a wide range of flowering plants (1-3). In contrast to our deep understanding of floral organs common to many angiosperms, genes that regulate the development of floral organs and/or flower-associated structures distinctive to a group of plant species remain poorly elucidated.Grass species, such as Oryza sativa (rice) and Zea mays (maize), bear a unique inflorescence consisting of spikelets and florets (4, 5). Each spikelet produces a defined number of florets depending on species, and is subtended by a pair of glumes. The floret comprises the flower proper (carpels, stamens, and lodicules) and a pair of additional structures (a palea and a lemma) that subtend the flower. The lodicule, an organ homologous to the petal in ordinary flowers, is small and semitransparent, and acts to open the palea and lemma for anthesis. Molecular genetic studies in rice and maize have revealed that the function of B class MADS-box genes is conserved in grasses: these genes specify the identities of the lodicule and stamen similar to the way in which B class genes specify t...