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First published online 27 February 2008
doi: 10.1242/dev.017913

Development 135, 1325-1334 (2008)
Published by The Company of Biologists 2008
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Ribosomal proteins promote leaf adaxial identity

Yao Yao*, Qihua Ling*, Hua Wang and Hai Huang{dagger}

National Laboratory of Plant Molecular Genetics, Shanghai Institute of Plant Physiology and Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, 300 Fenglin Road, Shanghai 200032, People's Republic of China.


Figure 1
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  		Fig. 1. Molecular identification of AE5 and AE6. (A,B) Fine structure mapping to localize AE5 and AE6 genes on chromosome 2 (A, upper panel) and 3 (B, upper panel), and AE5 (At2g19730) and AE6 (At3g25520) encode ribosomal proteins RPL28A (A, lower panel) and RPL5A (B, lower panel), respectively. Black and white boxes indicate the protein-coding region and untranslational region (UTR), respectively. (C) RT-PCR shows that AE5 and AE6 were expressed in all plant tissues examined. (D-J) In situ hybridization analyses to determine AE5 (D,F,H) and AE6 (E,G) expression patterns. (D) A late-globular-stage embryo. (E) A heart-stage embryo. (F) A longitudinal section of an 11-day-old seedling. (G) A transverse section of an 11-day-old seedling. (H) A longitudinal section of an inflorescence meristem. (I,J) No hybridization signals were detected either with the AE5 sense probe (I) or with the AE6 antisense probe for tissues from the insertional allele, ae6-2 (J). Blue lines in A,B indicate probe regions used for in situ hybridization. sam, shoot apical meristem; lp, leaf primordium; yl, young leaf; im, inflorescence meristem; fm, floral meristem; fp, flower primordium; s, sepal; st, stamen; c, carpel; an, anther. Scale bars: 25 µm in D,E; 100 µm in F-J.

 

Figure 2
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  		Fig. 2. ae5 and ae6 enhance as2 and as1 leaf polarity defects. (A-I) Phenotypes of wild-type and mutant seedlings in the Ler background. (A) Wild-type Ler. (B) as2-101, (C) as1-101. (D) ae5-1 as2-101. (E) ae6-1 as2-101. (F) ae5-1. (G) ae6-1. (H) ae5-1 as1-101. (I) ae6-1 as1-101. (J-M) Phenotypes of mutant seedlings in the Col-0 background. (J) ae5-2. (K) ae5-2 as2-1. (L) ae6-2. (M) ae6-2 as2-1. Arrows indicate the lotus-like leaves, and arrowheads indicate needle-like leaves. Scale bars: 0.5 cm.

 

Figure 3
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  		Fig. 3. Phenotypes of the ae5-1 and ae6-2 single mutants. (A,B) Primordia of the first two leaves in 3-day-old wild-type (A) and ae5-1 (B) seedlings. (C-F) Venations of the first-pair rosette leaves in wild-type Ler (C), wild-type Col-0 (D), ae5-1 (E) and ae6-2 (F). Insets in C-F show higher magnification of the vein pattern. (G-J) Transverse sections of the leaf blade in Ler (G), Col-0 (H), ae5-1 (I) and ae6-2 (J). Arrows indicate the normal abaxial intercellular spaces, while arrowheads show the ectopically positioned intercellular spaces. (K) An inflorescence of a wild-type Ler plant. (L-N) Inflorescences of ae5-1 showed various abnormalities, including disorganized inflorescences (L), abnormal phyllotaxy (M) and an ectopic leaf outgrowth that formed on distal part of the abaxial side of some cauline leaves (N, arrow). The arrowhead in M indicates two associated lateral branches; a cauline leaf was absent at the proximal end of one branch. Scale bars: 50 µm in A,B; 0.3 cm in C-F; 30 µm in G-J; 0.5 cm in K-N.

 

Figure 4
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  		Fig. 4. Aberrant adaxial-abaxial polarity in leaves of ae5-1 as2-101 and ae6-1 as2-101. (A-C) Three types of abnormal rosette leaves in ae5-1 as2-101. (A) An adaxial view of an expanded leaf showing a rough surface and some bumps (arrow). (B) A lotus-like leaf. (C) A needle-like leaf. (D-I) Analyses of the leaf epidermal patterns. (D,E) Epidermal cells on adaxial (D) and abaxial (E) surfaces of a wild-type leaf. (F,G) High magnification of epidermal cells in A. The same adaxial leaf side of ae5-1 as2-101 contained both adaxially (F) and abaxially (G) featured epidermal cells, which correspond to the boxed regions f and g in A, respectively. (H,I) High magnification of epidermal cells in C, corresponding to the boxed regions h and i, respectively. Arrowheads in E,G,I indicate long and narrow abaxially featured epidermal cells. (J-L) Transverse sections through the blade-petiole junction region. (J) A section from a wild-type Ler leaf. (K) A section from an expanded ae6-1 as2-101 leaf. (L) A section from an ae5-1 as2-101 lotus-like leaf. Arrowheads in J-L indicate phloem. ad, leaf adaxial side; ab, leaf abaxial side. Scale bars: 1 mm in A-C; 50 µm in D-L.

 

Figure 5
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  		Fig. 5. In situ localization of FIL and REV in ae5-1 and ae5-1 as2-101 mutants. (A-D) In situ hybridization of FIL in Ler (A), ae5-1 (B), as2-101 (C) and ae5-1 as2-101 (D). (E-H) In situ hybridization of REV in Ler (E), ae5-1 (F), as2-101 (G) and ae5-1 as2-101 (H). The FIL expression was adaxially extended in some primordia of as2-101 (C, arrowheads) and was apparent throughout some primordia of ae5-1 as2-101 (D, arrows). Arrowheads in H indicate vascular tissues.

 

Figure 6
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  		Fig. 6. Phenotypic analyses of double and triple mutants. (A) A rev-9 seedling. (B) A severe ae5-1 rev-9 seedling with only expanded cotyledons. An arrowhead indicates a needle-like leaf. (C) A less severe ae5-1 rev-9 seedling showing rosette leaves with rough leaf surface. (D) SEM of adaxial leaf surface of ae5-1 rev-9, showing that the abaxially featured epidermal cells appeared on the adaxial leaf side (arrowhead). (E) An rdr6-3 seedling. (F) An ae5-1 rdr6-3 seedling with lotus leaves (arrowheads). (G,H) Transverse sections of rdr6-3 (G) and ae5-1 rdr6-3 (H) leaves. The positions of sectioning are indicated by white lines in E and F. Arrowheads in G,H indicate the phloem. (I) SEM of a leaf from ett-3 arf4-2 double mutant, showing a number of outgrowths on the abaxial leaf side. (J) SEM of an ae5-1 ett-3 arf4-2 leaf, exhibiting a dramatically reduced number of the outgrowths. (K) A kan1-2 kan2-1 seedling. (L) An ae5-1 kan1-2 kan2-1 seedling. Insets in K,L show the abaxial surface of the first-pair leaf; arrowheads in J,K indicate the outgrowths. (M,N) Quantitative analyses of outgrowths on the third-pair (M) and fourth-pair (N) rosette leaves of ett-3 arf4-2 and ae5-1 ett-3 arf4-2. (O) Quantitative analysis of outgrowths on the first-pair rosette leaves of kan1-2 kan2-1 and ae5-1 kan1-2 kan2-1. Columns and error bars in M-O represent the mean and s.d. Scale bars: 0.5 cm in A-C,E,F; 50 µm in D,G,H; 3 mm in I,J; 2 mm in K,L; 1 mm in K,L insets.

 

Figure 7
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  		Fig. 7. Mutations in RPL24B and RPL5B enhance as2-1 polarity defects. (A) An stv1-1 seedling. (B) An stv1-1 as2-1 seedling. (C) SEM to examine epidermal cells on the top part of an stv1-1 as2-1 needle-like leaf. (D) Transverse section of an stv1-1 as2-1 needle-like leaf petiole. (E) An rpl5b seedling. (F) An rpl5b as2-1 seedling. (G) The top part of an rpl5b as2-1 needle-like leaf. (H) Transverse section of an rpl5b as2-1 needle-like leaf petiole. Arrowheads in C,G show the abaxially featured epidermal cells. Arrowheads in D,H indicate phloem. Scale bars: 0.5 cm in A,B,E,F; 50 µm in C,D,G,H.

 

Figure 8
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  		Fig. 8. Subcellular localization of YFP-RPL28A and YFP-RPL5A in the Arabidopsis root. (A-C) YFP-RPL28A was detected in both nuclei and cytoplasm. (D-I) Signals of YFP-RPL28A (D-F) and YFP-RPL5A (G-I) were found in both nucleolus and nucleoplasm. (A,D,G) YFP fluorescence; (B,E,H) bright-field differential interference contrast images; (C,F,I) merged views. Scale bars: 20 µm in A-C; 5 µm in D-I.

 

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© The Company of Biologists Ltd 2008