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First published online 15 September 2004
doi: 10.1242/dev.01388

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PIN-FORMED1 and PINOID regulate boundary formation and cotyledon development in Arabidopsis embryogenesis

Masahiko Furutani¹, Teva Vernoux^2,*, Jan Traas², Takehide Kato¹, Masao Tasaka¹ and Mitsuhiro Aida^1,

¹ Graduate School of Biological Sciences, Nara Institute of Science and Technology, Nara 630-0101, Japan
² INRA, Laboratoire de Biologie Cellulaire, Route de Saint Cyr, 78026 Versailles Cedex, France

View larger version (72K): [in a new window]   Fig. 1. Cotyledon development in wild type, pin1 and pid. (A-C) Five-day-old seedlings of wild type (A), pin1-3 (B) and pid-2 (C). (D-F) Scanning electron micrographs of wild-type (D), pin1-3 (E) and pid-2 (F) embryos. (G) ANT expression in wild-type embryo at the early heart stage, serial longitudinal sections. (H-K) FIL expression in wild-type (H), pin1-3 (I), pid-2 (J) and pin1-3 pid-2 (K) embryos at the early heart stage, serial longitudinal sections. Scale bars: 50 µm in D-K.

View larger version (133K): [in a new window]   Fig. 2. Phenotype of pin1 pid double mutant. (A-E) Five-day-old seedlings of pin1-3 pid-2 displaying severe (A,D), mild (B,E) and weak (C) phenotypes. (D,E) Scanning electron micrographs. Arrowheads in D indicate fused leaves with aberrant phyllotaxis. Scale bars: 100 µm in D, E.

View larger version (72K): [in a new window]   Fig. 3. PID mRNA expression in wild-type and pin1 embryos. (A-E) PID expression in wild-type embryos at the globular stage (A) and the early heart stage (B) in serial frontal longitudinal sections; the early heart stage in sagittal longitudinal section (C), and the torpedo stage in frontal (D) and sagittal (E) sections. (F,G) Serial longitudinal sections of pin1-3 embryo at the early heart stage (F) and the torpedo stage (G). (H) Schematic diagrams of PID expression in globular (left), heart (middle) and torpedo (right) stage embryos. Arrowheads in A indicate strong expression domains of PID mRNA. (C,E) Frontal view of each embryo with red line indicating the section plane. Scale bars: 50 µm in A-G.

View larger version (90K): [in a new window]   Fig. 4. Expression patterns of CUC1 and CUC2 in pid and pin1 pid at the early heart stage. CUC1 (A,C,E) and CUC2 (B,D,F) expression in serial longitudinal sections of wild-type (A,B), pid-2 (C,D) and pin1-3 pid-2 (E,F) embryos. Scale bars: 50 µm.

View larger version (124K): [in a new window]   Fig. 5. Relationship between PIN1, PID, CUC1, CUC2 and STM genes. (A-D) Expression of CUC genes in pin1-3 (A,B) and pin1-3 pid-2 (C,D) at the late heart stage. CUC1 expression (A,C) and CUC2 expression (B,D) in serial longitudinal sections. (E,F) STM expression in wild-type (E) and pin1-3 pid-2 (F) embryos at the heart stage in serial longitudinal sections. (G-J,O,P) Five-day-old seedlings of cuc1-1 cuc2 (G), pin1-3 pid-2 cuc1-1 (H), pin1-3 pid-2 cuc2 (I), pin1-3 pid-2 cuc1-1 cuc2 (J), stm-1 (O) and pin1-3 pid-2 stm-1 (P) plants. (K,L) Seven-day-old seedlings of cuc1-1 cuc2 (K) and pin1-3 pid-2 cuc1-1 cuc2 (L) plants were cleared to visualize the vascular pattern. (M,N) Vascular patterns of cuc1-1 cuc2 (M) and pin1-3 pid-2 cuc1-1 cuc2 (N) seedlings. Scale bars: 50 µm in A-F.

View larger version (110K): [in a new window]   Fig. 6. Effects of treatment with exogenous auxin on embryo. (A,B) Five-day-old seedlings treated with 2,4-D during embryogenesis. In B, the seedling was cleared to visualize the vascular pattern. (C,D) Spatial expression pattern of DR5::GUS in mature embryos. Mock (C) and 2,4-D (D) treatment. The fused part of cotyledons is indicated by a red arrowhead in D.

View larger version (74K): [in a new window]   Fig. 7. Apical patterning during embryogenesis. (A-D) Mutant phenotypes. (A,B) Apical region of the early-heart stage embryo. (A) Wild-type apex is divided into three subregions, presumptive SAM (PS; orange), cotyledon primordia (CP; yellow) and boundary of cotyledon margins (BCM; blue). (B) In the pin1-3 or pin1-3 pid-2 embryo, the peripheral region possesses a mixed identity of CP and BCM, which is shown in green. (C,D) Late heart stage. The area of BCM is reduced in pin1-3 (C), while the entire peripheral region continues to express a mixed identity of CP and BCM in pin1-3 pid-2 (D). (E,F) Model for apical patterning during embryogenesis. (E) PID expression (blue) accumulates mainly in the boundaries of cotyledon primordia and slightly in regions that surround the base of cotyledon primordia. PIN1 and PID redundantly promote auxin transport toward the tips of the cotyledon primordia (brown arrows), resulting in the formation of auxin gradient maxima (green). (F) Auxin accumulation in cotyledon primordia (green) prevents CUC gene expression (yellow) from expanding to the periphery.