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doi: 10.1242/10.1242/dev.00362

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HASTY, the Arabidopsis ortholog of exportin 5/MSN5, regulates phase change and morphogenesis

Krista M. Bollman, Milo J. Aukerman, Mee-Yeon Park, Christine Hunter, Tanya Z. Berardini and R. Scott Poethig*

Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018, USA



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  		Fig. 1. The phenotype of hst-1. (A) Two-week-old wild-type rosette (B) Two-week-old hst-1 rosette. (C) Wild-type inflorescence. (D) hst-1 inflorescence. Note abnormally expanded internodes and short lateral inflorescences (arrow) and unfertilized siliques (asterisk). (E) Wild-type flowers. (F) hst-1 flowers. hst-1 flowers are unusually small and are irregularly spaced (arrow).

 


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  		Fig. 2. The effect of hst on seedling morphology. (A) Three-day-old wild-type (left) and hst-1 (right) seedlings grown in the dark. (B) Seven-day-old wild-type (left) and hst-1 (right) seedlings grown in the light. (C-E) Anatomy of 3-day-old wild-type seedlings. (F-H) Anatomy of 3-day-old hst-1 seedlings. (C,F) Median longitudinal section of the shoot apex; the meristem is marked with an asterisk. (D,G) Cross-section of the hypocotyl. (E,H) Cross-section of the root. (I) Length of the hypocotyl and root in dark- and light-grown seedlings. Black, wild-type hypocotyl; white, wild-type root; white hatching on black, hst-1 hypocotyl; black hatching on white, hst-1 root. Scale bars: 50 µm.

 


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  		Fig. 3. Effect of hst-1 on phase change. (A) Rosette leaf and bract morphology of hst-1 and wild-type plants (Col). Gray, no abaxial trichomes; black, abaxial trichomes; underline, bracts. (B) Adaxial trichome density of rosette leaves and bracts in hst-1 and wild-type plants (C) The blade:petiole length ratio of rosette leaves in hst-1 and wild-type plants. (D) Rate of leaf initiation in hst-1 and wild-type plants. Arrows indicate the position of the first leaf with abaxial trichomes in siblings of the dissected plants; asterisks indicate the time at which GUS-stained floral buds were visible. White circles, wild type; black circles, hst-1.

 


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  		Fig. 4. Effect of hst-1 on leaf and carpel polarity. (A,B) Paradermal views of the adaxial and abaxial mesophyll layers of leaf 3 from wild-type (A) and hst-1 (B) plants. (C-E) Scanning electron micrographs of wild-type (C) and hst-1 (D,E) pistils; note the ovules located along the margins of the carpels (arrow). (F) Phenotype of kan-11, hst-1 and hst-1; kan-11 seedlings. Double mutant seedlings have more severely up-rolled leaves and cotyledons, and have a significantly larger number of abaxial trichomes than either single mutant.

 


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  		Fig. 5. The genetic interaction between hst-1 and sqn-1. Double mutants are smaller and produce many more abaxial trichomes on leaves 1 and 2 than single-mutant plants.

 


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  		Fig. 6. Positional cloning of HST. The top line indicates the genetic map of the region surrounding hst-1, as determined from an analysis of F2 progeny from a cross to Ler. The number of recombinants in each interval is indicated below the line. BACs that hybridized to the lamba clone m262, which showed no recombination with hst-1, are shown below this line. Hybridization of F9J1 to an EcoRV digest of genomic DNA from hst-9 and Col revealed a missing fragment in hst-9. When this EcoRV fragment was gel purified from F9L1 DNA and used to re-probe the blot, it hybridized to two fragments in hst-9. These fragments are not visible in the blot hybridized with the entire BAC because this blot was under-exposed in order to visualize as many fragments as possible.

 


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  		Fig. 7. Molecular characterization of HST. (A) Genomic structure. Exons are indicated by boxes. Arrows indicate the position of mutations described in the text. (B) Poly(A)-enriched RNA from leaves, roots and floral buds hybridized with a 5' fragment of the HST cDNA. This blot was stripped and re-hybridized with an actin probe, which served as a loading control. (C) CLUSTAL X 1.81 alignment of the amino acid sequence of HST (AY198396), human Xpo5 (NP_065801) and Msn5p (NP_010622) from S. cerevisiae. Amino acids identical in two or more sequences are highlighted in black; similar amino acids are highlighted in gray.

 


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  		Fig. 8. Unrooted bootstrap (1000 reiterations) cladogram of human and Arabidopsis importin ß-like proteins. This tree was generated by neighbor-joining using Clustal X 1.81 and was visualized using TreeView. Black, human; blue, Arabidopsis; red, HST. Accession Numbers for human proteins: CAS (NP_001307), KAPß3 (NP_002262), IMPß (NP_002256), IMP4 (NP_078934), IMP9 (NP_060555), IMP11 (NP_057422), IMP13 (NP_055467), RANBP6 (AAC14260), RANBP7 (NP_006382), RANBP8 (NP_006381), RANBP16 (NP_055839), RANBP17 (NP_075048), TRN (NP_002261), TRN2 (NP_038461), TRNSR (NP_036602), XPO1 (NP_003391), XPO4 (BAB21812), XPO5 (NP_065801), XPOT (NP_009166). Sequences of Arabidopsis proteins are available at www.tigr.org/tdb/e2k1/ath1/LocusNameSearch.shtml.

 


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  		Fig. 9. Two hybrid analysis of the interaction between HST and RAN1. Cells transformed simultaneously with the indicated vectors were selected for the presence of both constructs on medium lacking Leu and Trp, and then plated on a medium lacking Leu, Trp, His and Ade to test for the interaction between the AD and BD constructs. Cells containing the HST:BD and RAN1:AD vectors grew on this restrictive medium and expressed ß galactosidase (not shown). Cells containing RAN1:AD and either the hst-3:BD or {Delta}N-HST:BD vectors grew very poorly or not at all on this medium, and did not express ß galactosidase.

 


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  		Fig. 10. Location of GUS activity in hst-1 plants homozygous for a 35S::HST-GUS transgene. (A) Seedling root, (B) trichome and (C) trichome of a plant transformed with the 35S::GUS vector used in the production of the 35S::HST-GUS fusion construct.

 

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© The Company of Biologists Ltd 2003