The complex of <scp>ASYMMETRIC LEAVES</scp> (<scp>AS</scp>) proteins plays a central role in antagonistic interactions of genes for leaf polarity specification in Arabidopsis

Machida, Chiyoko; Nakagawa, Ayami; Kojima, Shoko; Takahashi, Hiro; Machida, Yasunori

doi:10.1002/wdev.196

Cited by 65 publications

(83 citation statements)

References 113 publications

(167 reference statements)

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“…Since the frequencies of formation of filamentous leaves were increased in the as1 or as2 mutant treated with hydroxyurea or in the double mutants, as1-1 icu2-1, as2-1 icu2-1, as1-1 rfc3-1, as2-1 rfc3-1, the AS1-AS2-mediated pathway and the progression of DNA replication might be independently, but cooperatively, involved in the development of the adaxial domain of leaves in the wildtype Arabidopsis ( Figure 5). Such cooperative actions in the leaf development are also observed in combinations of as mutations with other modifiers (Machida et al 2015).…”

Section: Discussionmentioning

confidence: 89%

“…AS1 and AS2 encode nuclear proteins and form a complex (designated AS1-AS2) (Guo et al 2008;Luo et al 2012;Xu et al 2003;Yang et al 2008). Mutations in these genes are associated with pleiotropic abnormalities in leaves along the three developmental axes described above (Byrne et al 2000;Iwakawa et al 2002Matsumura et al 2009;Ori et al 2000;Rédei and Hirono 1964;Semiarti et al 2001;Tsukaya and Uchimiya 1997), suggesting that AS1-AS2 regulates multiple genes Machida et al 2015;Takahashi et al 2008;Takahashi et al 2013) that might be involved in leaf formation along the three axes.…”

Section: Introductionmentioning

confidence: 99%

“…Defects in polarity of as1 and as2 leaves are enhanced under certain growth conditions as well as in conjunction with mutations in members of certain groups of genes (Machida et al 2015), which are designated as modifiers of adaxial-abaxial patterning (Machida et al 2015;Matsumura et al 2016;Szakonyi et al 2010). To date, various modifier mutations that generate filamentous leaves surrounded by abaxialized epidermis in the as1 or as2 mutant backgrounds have been identified and it has been reported that the development of the adaxial domain of leaves is severely defective in those double mutants.…”

Section: Introductionmentioning

confidence: 99%

“…To date, various modifier mutations that generate filamentous leaves surrounded by abaxialized epidermis in the as1 or as2 mutant backgrounds have been identified and it has been reported that the development of the adaxial domain of leaves is severely defective in those double mutants. The modifier genes include several that mediate the biogenesis of tasiR-ARF (Machida et al 2015;Yang et al 2006). Other relevant modifier genes belong to several different groups: those for ribosome biogenesis (Horiguchi et al 2011a;Matsumura et al 2016;Pinon et al 2008;Yao et al 2008); chromatin modification (Kojima et al 2011;Ueno et al 2007); and cell proliferation (Horiguchi et al 2011b;Ishibashi et al 2012;Wang et al 2011;Xu et al 2012;Yuan et al 2010).…”

Section: Introductionmentioning

confidence: 99%

“…Other relevant modifier genes belong to several different groups: those for ribosome biogenesis (Horiguchi et al 2011a;Matsumura et al 2016;Pinon et al 2008;Yao et al 2008); chromatin modification (Kojima et al 2011;Ueno et al 2007); and cell proliferation (Horiguchi et al 2011b;Ishibashi et al 2012;Wang et al 2011;Xu et al 2012;Yuan et al 2010). Mutations of genes related to DNA replicationrelated chromatin remodeling and DNA repair have been identified as modifiers that enhance leaf adaxialabaxial abnormalities in as1 and as2 (Inagaki et al 2009;Machida et al 2015;Xu et al 2012). Mutations in FASCIATA1 (FAS1) and FAS2 genes that encode Arabidopsis homologs of components of Chromatin Assembly Factor-1 (CAF-1), a histone chaperone essential for chromatin remodeling after DNA replication (Takami et al 2007), also cause abaxialized filamentous leaves in as1 and as2 Xu et al 2012).…”

Section: Introductionmentioning

confidence: 99%

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A genetic link between epigenetic repressor AS1–AS2 and DNA replication factors in establishment of adaxial–abaxial leaf polarity of Arabidopsis

Luong

Keta

Asai

et al. 2018

Plant Biotechnology

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Balanced development of adaxial and abaxial domains in leaf primordia is critical for the formation of flat symmetric leaf lamina. Arabidopsis ASYMMETRIC LEAVES1 (AS1) and AS2 proteins form a complex (AS1-AS2), which acts as key regulators for the adaxial development by the direct repression of expression of the abaxial gene ETTIN/AUXIN RESPONSE FACTOR3 (ETT/ARF3). Many modifier mutations have been identified, which enhance the defect of as1 and as2 mutations to generate abaxialized filamentous leaves without adaxial traits, suggesting that the development of the adaxial domain is achieved by cooperative repression by AS1-AS2 and the wild-type proteins corresponding to the modifiers. Mutations of several genes for DNA replication-related chromatin remodeling factors such as Chromatin Assembly Factor-1 (CAF-1) have been also identified as modifiers. It is still unknown, however, whether mutations in genes involved in DNA replication themselves might act as modifiers. Here we report that as1 and as2 mutants grown in the presence of hydroxyurea, a known inhibitor of DNA replication, form abaxialized filamentous leaves in a concentration-dependent manner. We further show that a mutation of the INCURVATA2 (ICU2) gene, which encodes the putative catalytic subunit of DNA polymerase α, and a mutation of the Replication Factor C Subunit3 (RFC3) gene, which encodes a protein used in replication as a clamp loader, act as modifiers. In addition, as2-1 icu2-1 double mutants showed increased mRNA levels of the genes for leaf abaxialization. These results suggest a tight link between DNA replication and the function of AS1-AS2 in the development of flat leaves.

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Section: Discussionmentioning

confidence: 89%

Section: Introductionmentioning

confidence: 99%

Section: Introductionmentioning

confidence: 99%

Section: Introductionmentioning

confidence: 99%

Section: Introductionmentioning

confidence: 99%

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A genetic link between epigenetic repressor AS1–AS2 and DNA replication factors in establishment of adaxial–abaxial leaf polarity of Arabidopsis

Luong

Keta

Asai

et al. 2018

Plant Biotechnology

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Peptide signaling molecules CLE5 and CLE6 affect Arabidopsis leaf shape downstream of leaf patterning transcription factors and auxin

et al. 2018

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Intercellular signaling mediated by small peptides is critical to coordinate organ formation in animals, but whether extracellular polypeptides play similar roles in plants is unknown. Here we describe a role in Arabidopsis leaf development for two members of the CLAVATA 3/ ESR ‐ RELATED peptide family, CLE 5 and CLE 6, which lie adjacent to each other on chromosome 2. Uniquely among the CLE genes, CLE 5 and CLE 6 are expressed specifically at the base of developing leaves and floral organs, adjacent to the boundary with the shoot apical meristem. During vegetative development CLE 5 and CLE 6 transcription is regulated by the leaf patterning transcription factors BLADE ‐ ON ‐ PETIOLE 1 ( BOP 1) and ASYMMETRIC LEAVES 2 ( AS 2), as well as by the WUSCHEL ‐ RELATED HOMEOBOX ( WOX ) transcription factors WOX 1 and PRESSED FLOWER ( PRS ). Moreover, CLE 5 and CLE 6 transcript levels are differentially regulated in various genetic backgrounds by the phytohormone auxin. Analysis of loss‐of‐function mutations generated by genome engineering reveals that CLE 5 and CLE 6 independently and together have subtle effects on rosette leaf shape. Our study indicates that the CLE 5 and CLE 6 peptides function downstream of leaf patterning factors and phytohormones to modulate the final leaf morphology.

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Arabidopsis ASYMMETRIC LEAVES2 (AS2): roles in plant morphogenesis, cell division, and pathogenesis

et al. 2021

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The ASYMMETRIC LEAVES2 (AS2) gene in Arabidopsis thaliana is responsible for the development of flat, symmetric, and extended leaf laminae and their vein systems. AS2 protein is a member of the plant-specific AS2/LOB protein family, which includes 42 members comprising the conserved amino-terminal domain referred to as the AS2/LOB domain, and the variable carboxyl-terminal region. Among the members, AS2 has been most intensively investigated on both genetic and molecular levels. AS2 forms a complex with the myb protein AS1, and is involved in epigenetic repression of the abaxial genes ETTIN/AUXIN RESPONSE FACTOR3 (ETT/ARF3), ARF4, and class 1 KNOX homeobox genes. The repressed expression of these genes by AS2 is markedly enhanced by the cooperative action of various modifier genes, some of which encode nucleolar proteins. Further downstream, progression of the cell division cycle in the developing organs is stimulated; meristematic states are suppressed in determinate leaf primordia; and the extension of leaf primordia is induced. AS2 binds the specific sequence in exon 1 of ETT/ARF3 and maintains methylated CpGs in several exons of ETT/ARF3. AS2 forms bodies (designated as AS2 bodies) at nucleolar peripheries. AS2 bodies partially overlap chromocenters, including inactive 45S ribosomal DNA repeats, suggesting the presence of molecular and functional links among AS2, the 45S rDNAs, and the nucleolus to exert the repressive regulation of ETT/ARF3. The AS2/LOB domain is characterized by three subdomains, the zinc finger (ZF) motif, the internally conserved-glycine containing (ICG) region, and the leucine-zipper-like (LZL) region. Each of these subdomains is essential for the formation of AS2 bodies. ICG to LZL are required for nuclear localization, but ZF is not. LZL intrinsically has the potential to be exported to the cytoplasm. In addition to its nuclear function, it has been reported that AS2 plays a positive role in geminivirus infection: its protein BV1 stimulates the expression of AS2 and recruits AS2 to the cytoplasm, which enhances virus infectivity by suppression of cytoplasmic post transcriptional gene silencing.

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The complex of ASYMMETRIC LEAVES (AS) proteins plays a central role in antagonistic interactions of genes for leaf polarity specification in Arabidopsis

Cited by 65 publications

References 113 publications

A genetic link between epigenetic repressor AS1–AS2 and DNA replication factors in establishment of adaxial–abaxial leaf polarity of <i>Arabidopsis</i>

A genetic link between epigenetic repressor AS1–AS2 and DNA replication factors in establishment of adaxial–abaxial leaf polarity of <i>Arabidopsis</i>

Peptide signaling molecules CLE5 and CLE6 affect Arabidopsis leaf shape downstream of leaf patterning transcription factors and auxin

Arabidopsis ASYMMETRIC LEAVES2 (AS2): roles in plant morphogenesis, cell division, and pathogenesis

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