title: The complex of ASYMMETRIC LEAVES (AS) proteins plays a central role in antagonistic interactions of genes for leaf polarity specification in Arabidopsis.
abstract: Leaf primordia are born around meristem-containing stem cells at shoot apices, grow along three axes (proximal-distal, adaxial-abaxial, medial-lateral), and develop into flat symmetric leaves with adaxial-abaxial polarity. Axis development and polarity specification of Arabidopsis leaves require a network of genes for transcription factor-like proteins and small RNAs. Here, we summarize present understandings of adaxial-specific genes, ASYMMETRIC LEAVES1 (AS1) and AS2. Their complex (AS1-AS2) functions in the regulation of the proximal-distal leaf length by directly repressing class 1 KNOX homeobox genes (BP, KNAT2) that are expressed in the meristem periphery below leaf primordia. Adaxial-abaxial polarity specification involves antagonistic interaction of adaxial and abaxial genes including AS1 and AS2 for the development of two respective domains. AS1-AS2 directly represses the abaxial gene ETTIN/AUXIN RESPONSE FACTOR3 (ETT/ARF3) and indirectly represses ETT/ARF3 and ARF4 through tasiR-ARF. Modifier mutations have been identified that abolish adaxialization and enhance the defect in the proximal-distal patterning in as1 and as2. AS1-AS2 and its modifiers synergistically repress both ARFs and class 1 KNOXs. Repression of ARFs is critical for establishing adaxial-abaxial polarity. On the other hand, abaxial factors KANADI1 (KAN1) and KAN2 directly repress AS2 expression. These data delineate a molecular framework for antagonistic gene interactions among adaxial factors, AS1, AS2, and their modifiers, and the abaxial factors ARFs as key regulators in the establishment of adaxial-abaxial polarity. Possible AS1-AS2 epigenetic repression and activities downstream of ARFs are discussed.
authors.affiliation: Graduate School of Bioscience and Biotechnology, Chubu University, Kasugai, Japan.
authors.forename: Chiyoko
authors.initials: C
authors.lastname: Machida
authors.affiliation: Graduate School of Bioscience and Biotechnology, Chubu University, Kasugai, Japan.
authors.forename: Ayami
authors.initials: A
authors.lastname: Nakagawa
authors.affiliation: Graduate School of Bioscience and Biotechnology, Chubu University, Kasugai, Japan.
authors.forename: Shoko
authors.initials: S
authors.lastname: Kojima
authors.affiliation: Graduate School of Horticulture, Chiba University, Chiba, Japan.
authors.forename: Hiro
authors.initials: H
authors.lastname: Takahashi
authors.affiliation: Graduate School of Science, Nagoya University, Nagoya, Japan.
authors.forename: Yasunori
authors.initials: Y
authors.lastname: Machida
mesh_terms: D017360:Arabidopsis / Q000235:genetics* / Q000254:growth & development*; D029681:Arabidopsis Proteins / Q000235:genetics*; D019521:Body Patterning / Q000235:genetics*; D018506:Gene Expression Regulation, Plant / Q000235:genetics; D009154:Mutation / Q000235:genetics; D018515:Plant Leaves / Q000196:embryology* / Q000235:genetics*; D014157:Transcription Factors / Q000235:genetics
chemical_list: D029681:Arabidopsis Proteins; D014157:Transcription Factors