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doi: 10.1242/10.1242/dev.00588

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The Drosophila bZIP transcription factor Vrille is involved in hair and cell growth

Sébastien Szuplewski*, Benjamin Kottler and Régine Terracol{dagger}

Laboratoire de Génétique du Développement et Evolution, Institut Jacques Monod, 2 Place Jussieu Tour 43, 75251 Paris Cedex 05, France



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  		Fig. 1. Mapping of vri 5' end and generation of deletion alleles by P-element mutagenesis. The position of the vri alleles and the extent of the deletions generated are shown. The localization of the vri cDNAs, the other described loci, the putative GadFly genes and the BDGP cDNAs are presented. The positions of the genomic (1, 2, 3, 4) and PlacW primers used for PCR amplification are shown (arrows). vri1 (cDNA3.8) and vri2 (cDNA3.3) are the two types of cDNAs used for UAS constructs. DS00714 distal end of a P1 phage isolated by the BDGP, DM194B8, T7 end and DM77B6, SP6 end of two cosmids isolated by the European Drosophila Mapping Consortium.

 


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  		Fig. 2. vri affects wing shape and hair morphology and interacts with actin-binding protein encoding genes. (A) Wild-type adult female fly. (B) vri11/vri5 adult female fly with shorter downward bending wings and shorter and upturned posterior scutellar bristles (arrow). (C) Wild-type wing. (D) vri11/vri5R5.24 wing, smaller than wild type with notching at the anterior margin (right-hand arrow) and multiple regions with atrophic or missing hairs appearing as non pigmented zones on the wing surface (arrows). (E) vri5R7.2/kst01318 wing showing missing hairs at the anterior margin (top arrow) and notching at the posterior margin (bottom arrow). (F) vri5R7.2/ActnG0077 wing with missing hairs at the anterior margin (top arrow) and on the wing surface (bottom arrow).

 


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  		Fig. 3. Mutations in vri decrease hair and cell size in adult cuticle. y/y; vri/vri somatic clones were induced at third larval instar by the FLP/FRT technique and observed in females. (A) y/y; vri2/vri2 clonal left posterior scutellar macrochaete (arrow) with a reduced size compared with the y+/y; vri+/vri right one. (B) vri1 clone showing a socket with no macrochaete on the head (arrow). (C) vri5 clones with pigmented cuticle. (D) y/y; vri5/vri5 clones at the wing anterior margin in the triple row region with smaller and thinner stout mechanosensory bristles. The space between bristles is reduced (vertical arrows), which is characteristic of smaller cells, and the margin is concave (double-headed arrow). (E) At the anterior wing margin, the average distance between stout bristles of the triple row is significantly reduced in y/y; vri/vri clones (Student's test: P<0.001) compared with y/y; vri+/vri+ clones. The values are percentages of wild type±s.e. Clones, analyzed from 12 to 31 wings, contained from two to 33 bristles.

 


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  		Fig. 4. vri loss of function allele reduces eye size, alters morphology and size of ommatidia and photoreceptors. Scanning electron micrographs of vri clonal eye were generated by the EGUF/hid method. Photoreceptor cells bearing the dominant photoreceptor cell lethal transgene GMR-hid die during metamorphosis. The l(2)CL-L1 recessive cell lethal mutation leads to lethality at an earlier stage of eye development. The crosses were performed at 29°C and the eyes were observed in males. (A,C,F) vri+ FRT40A/vri+ FRT40A control eyes from [ry+ hs-neo FRT]40A/[ry hs-neo FRT]40A GMR-hid l(2)CL-L1; ey-GAL4 UAS-FLP/+ flies. (B,D,G) vri5R7.2/vri5R7.2 clonal eyes from vri5R7.2[ry+ hs-neo FRT]40A/[ry hs-neo FRT]40A GMR-hid l(2)CL-L1; ey-GAL4 UAS-FLP/+ flies. (B) Clonal eye smaller than control (A) and with a rough aspect. (C,D) The ommatidia are disorganized and present an abnormal morphology. They are spherical rather than hexagonal and bristles are missing or atrophic. (E) The number of ommatidia is reduced in vri clonal eyes. Values are mean numbers of ommatidia (±s.e.) observed in male eyes. The mean number of ommatidia is reduced by 10% in vri5, 17% in vri1 and 19% in vri5R7.2 eyes compared with the vri+ control eyes (Student's test: P<0.001). (F,G) Longitudinal sections through adult retinas. (G) Clonal retina with photoreceptor stalks twisted and shorter than control (F).

 


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  		Fig. 5. vri overexpression induces atrophy of embryonic epidermis, adult wing and thoracic hairs. (A) Wild-type embryo. (B) UAS-vri16/pnr-GAL4 dead embryo showing atrophic dorsal epidermis and anterior hole and head defects. (C) pnr-GAL4/UAS-rbf dead embryo with a similar phenotype. (D) Wild-type wing. (E) vg-GAL4/+; UAS-vri113/+ wing at 29°C showing notching at the margin. (F) vg-GAL4/+; UAS-rbf/+ wing at 25°C with notching of the posterior margin. (G) Scanning electron micrographs of wild-type thorax. (H) hsp70-GAL4/UAS-vri19 thorax at 29°C showing atrophic macrochaete. (I) hsp70-GAL4/UAS-rbf thorax at 29°C with a similar phenotype.

 


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  		Fig. 6. vri overexpression induces atrophy of the eye. Scanning electron micrographs of ey-GAL4/UAS-vri19 eye series with progressive reduction of the eye at 25°C (A,E), and extreme phenotype with UAS-vri114/+; ey-GAL4/UAS-vri113 at 29°C (F). (A) Rough eye with a moderate reduction in size. (B,C) Ommatidia are disorganized with a globular aspect, bristles are duplicated or missing and regions of apparent irregular growth are observed (B). (D) Very reduced eye. (E) Eye totally absent. (F) At 29°C the UAS-vri114/+; ey-GAL4/UAS-vri113 flies lack all head structures derived from eye-antennal disc. The proboscis, which is derived from the labial disc, is present.

 


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  		Fig. 7. vri overexpression alters trichome number, size and morphology in wing. All wings are from females. (A) A9-GAL4/+ control wing. (B) Magnified view of A9-GAL4/+ control wing around L4 vein. (C) A9-GAL4/UAS-vri114; UAS-vri113/+ wing with a slightly reduced size, enlarged cells and reduced L5 vein. Long ectopic bristles are observed (magnification in the top right-hand corner). (D) Magnified A9-GAL4/UAS-vri114; UAS-vri113/+ wing surface, around L4 vein. Larger cells are observed with multiple trichomes. (E) MS1096-GAL4/+ control wing. (F) Magnified view of MS1096-GAL4/+ control wing around L3 vein. (G) MS1096-GAL4/+; UAS-vri28/+ wing. (H) Magnified MS1096-GAL4/+; UAS-vri28/+ wing showing enlarged L3 vein, enlarged cell size with multiple trichomes, some of them with an abnormal morphology. (I) MS1096-GAL4/+; UAS-vri28/UAS-vri28 wing with a greatly reduced size and an abnormal morphology. (J) Magnified MS1096-GAL4/+; UAS-vri28/UAS-vri28 vein around L3 vein showing multiple trichomes with a reduced size and an abnormal differentiation.

 


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  		Fig. 8. vri overexpression induces apoptosis and inhibits cell growth in larval tissues. Tissues were dissected at third larval instar. (A,B) Wing imaginal discs stained with Acridine Orange. (A) MS1096-GAL4/+ control disc with low level of apoptosis. (B) MS1096-GAL4/+; MS1096-GAL4/+; UAS-vri19/+ with increased number of apoptotic cells (arrow) in the wing pouch region. (C,D) DAPI staining of male salivary gland. (C) F4-GAL4/+ control gland. (D) F4-GAL4/+; UAS-vri19/+ gland (same magnification as control) with a size reduced by about one half and with pyknotic nuclei (arrows). (E,F) Fat body clones co-expressing vri and GFP induced by the `flip out' technique in hsp FLP; UAS-vri19/Act5C<CD2<GAL4, UAS-GFP females. (E) GFP detection (green) with vri overexpressing clones of greater green intensity. (F) GFP and DAPI (red) stainings showing clones (yellow) with smaller cells.

 

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© The Company of Biologists Ltd 2003