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First published online 27 February 2008
doi: 10.1242/dev.016469

Development 135, 1315-1324 (2008)
Published by The Company of Biologists 2008
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Three PIGGYBACK genes that specifically influence leaf patterning encode ribosomal proteins

Violaine Pinon1,*, J. Peter Etchells1,*,{dagger}, Pascale Rossignol1, Sarah A. Collier1, Juana M. Arroyo2, Robert A. Martienssen2 and Mary E. Byrne1,{ddagger}

1 Department of Crop Genetics, John Innes Centre, Norwich NR4 7UH, UK.
2 Cold Spring Harbor Laboratory, Cold Spring Harbor, NY 11724, USA.


Figure 1
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  		Fig. 1. Arabidopsis pgy1, pgy2 and pgy3 leaf phenotypes. (A) Rosettes of pgy mutants. From left to right: wild type, pgy1, pgy2, pgy3. (B) Rosettes of pgy mutants in as1 background. From left to right: as1, as1 pgy1, as1 pgy2, as1 pgy3. (C) Rosette leaves with ectopic outgrowths. From left to right: as1 pgy1, as1 pgy2, as1 pgy3. (D) Rosette of as1 pgy1 pgy2 pgy3 quadruple mutant.

 

Figure 2
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  		Fig. 2 . Development of ectopic outgrowths in Arabidopsis as1 pgy1. (A-C) Adaxial surface of a late rosette leaf from 12-day-old plants of wild type (A), as1 (B) and as1 pgy1, showing a dome-shaped structure (arrow) of an initiating outgrowth (C). (D,E) as1 pgy1 late rosette leaves from 15-day-old plants, showing ectopic outgrowths forming peg-like structures. (F) as1 pgy1 late rosette leaf from an 18-day-old plant forms a leaf-like structure with trichomes on the adaxial side but not on the abaxial side of an ectopic outgrowth. Scale bars: 100 µm.

 

Figure 3
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  		Fig. 3. Vascular patterning defect in Arabidopsis as1 pgy1 leaves. Cross sections through leaf midveins. In wild type (A), as1 (B) and pgy1 (C), xylem is adaxial and phloem is abaxial. In as1 pgy1 mutants (D), xylem is surrounded by phloem. ph, phloem; x, xylem. Scale bars: 50 µm.

 

Figure 4
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  		Fig. 4. rev enhances as1 and as1 pgy1 leaf phenotypes in Arabidopsis. (A) rev, with narrow and elongated leaves. (B) pgy1 rev, with slightly narrower leaves than rev. (C) as1. (D) as1 rev, exhibiting narrower and a more rippled leaf surface compared with as1. (E) as1 pgy1 rev triple mutants have a more severe phenotype than as1 pgy1 double mutants, with formation of very small rosettes accumulating anthocyanin in most leaves. (F) SEM of as1 pgy1 rev rosette, showing radial leaves (arrow). Scale bar: 50 µm in F.

 

Figure 5
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  		Fig. 5. pgy1 enhances rev axillary meristem defects in Arabidopsis. (A) Wild-type flowering plant. (B) rev mutants lack axillary meristems at the base of some cauline leaves (arrows). (C) pgy1 rev plant with reduced stature compared with wild type and rev. Inflorescence forms few or no flowers. (D) SEM of pgy1 rev inflorescence. Most floral meristems either produce filamentous structures or fail to produce any organs. (E) rev phb phv/+ plant with reduced stature and inflorescence with few flowers. (F) SEM of rev phb phv/+ inflorescence. Most floral meristems produce filamentous structures similar to those in pgy1 rev. Scale bars: 200 µm in D,F.

 

Figure 6
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  		Fig. 6. kan1 suppresses the as1 pgy1 leaf phenotype in Arabidopsis. (A-C) Rosettes of as1 (A), as1 kan1 (B) and as1 kan1 kan2/+ (C). The rosette leaves of as1 kan1 (B) and as1 kan1 kan2/+ (C) have a surface slightly less rippled than as1 (A). (D-F) Rosettes of as1 pgy1 (D), as1 pgy1 kan1 (E) and as1 pgy1 kan1 kan2/+ (F). The as1 pgy1 phenotype is suppressed by kan1, and to a greater extent in a kan1 kan2/+ background, such that as1 pgy1 kan1 kan2/+ rosette leaves are more similar to as1. (G-J) Plants of kan1 kan2 (G), as1 kan1 kan2 (H), pgy1 kan1 kan2 (I) and as1 pgy1 kan1 kan2 (J). The kan1 kan2 phenotype is not suppressed by as1, pgy1, or by as1 pgy1. (K) Quantitative RT-PCR was used to assay expression levels of PHB, PHV, REV, KAN1 and KAN2 in the following genotypes: wild type, pgy1, as1 and as1 pgy1. Results were normalised to ACTIN2, with the value from wild-type plants arbitrarily set to 1.0. Bars indicate the standard deviation between three biological replicates. ANOVA statistical analysis indicates a significant difference between Ler and as1 pgy1 for KAN1 and KAN2 expression. Scale bars: 1 cm in G-J.

 

Figure 7
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  		Fig. 7. pgy1 suppresses the kan1 kan2/+ gynoecium phenotype in Arabidopsis. (A-D) Flowering inflorescences of wild type (A), kan1 kan2/+ (B), as1 kan1 kan2/+ (C) and pgy1 kan1 kan2/+ (D). (E-H) Single siliques of wild type (E), kan1 kan2/+ (F), as1 kan1 kan2/+ (G) and pgy1 kan1 kan2/+ (H). kan1 kan2/+ and as1 kan1 kan2/+ siliques exhibit a similar phenotype, with proliferation of septum and formation of ectopic styles along the abaxial replum, between the two fused carpels (arrows). pgy1 kan1 kan2/+ silique (H) has less ectopic tissue and is more elongated compared with kan1 kan2/+ siliques (F). Scale bars: 1 mm in E-H.

 

Figure 8
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  		Fig. 8. Arabidopsis PGY genes encode ribosomal proteins. (A) Schematic representation of PGY1/RPL10aB, PGY2/RPL9C, PGY3/RPL5A genes showing sites of point mutations in pgy1-1, pgy1-2, pgy2-1, pgy3-1 alleles and site of T-DNA insertion in pgy2-2. (B-D) mRNA expression patterns of PGY1 (B), PGY2 (C) and PGY3 (D) in transverse sections of 10-day-old vegetative apices. All three genes are expressed in the meristem and throughout leaf primordia. In older leaves expression is higher in the vasculature and at the margins of the leaf, in cytoplasmically dense cells. Scale bars: 50 µm.

 

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