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First published online 4 April 2007
doi: 10.1242/dev.02844

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tailup, a LIM-HD gene, and Iro-C cooperate in Drosophila dorsal mesothorax specification

Centro de Biología Molecular Severo Ochoa, CSIC and UAM, Cantoblanco, 28049 Madrid, Spain.

View larger version (70K): [in this window] [in a new window]   Fig. 1. Cuticular phenotypes associated with tup clones. (A) View of the lateral-posterior region of a notum displaying ectopic structures in a fly bearing f tup2 clones. The framed area is shown at high magnification in B, after mounting of the cuticle. (B) S1 and S2, extant first and second axilary sclerites; S1' and S2', recognisable ectopic sclerites; asterisk and tg', mass of sclerotised tissue and an ectopic tegula-like structure, respectively, bearing macrochaetae and sensilla trichoidea. (C) Ectopic tegula (arrow) with ytupex4 bristles, and extant tegula (arrowhead). (D) Pedicel of a haltere (asterisk) showing rows of sensilla campaniformia (white arrowheads). Black arrowhead, ectopic structure on the metanotum bearing at its base pedicel-like sensilla campaniformia (arrows). Insets show boxed regions at high magnification. (E) Protuberance on the anterior lateral notum with f tup2 microchaetae (arrowhead). (F) Notum showing loss of tissue and of bristles in the scutellum (arrow) and a vesicular invagination (arrowhead). (G) High magnification of the vesicle shown in F. Arrowhead, group of sensilla campaniformia. (H,I) Two focal planes of an invagination with y tupex4 microchaetae arising from its interior (arrowhead). Green line, contour of a crinkled (ck)-marked twin-spot which covers the invagination. (J) Cuticular defects (arrow and arrowheads) on the scutum and scutellum of a fly with ftupisl-1 clones. (K) Protuberance/invagination (indicated by the arrow in J) at high magnification. Note the apical hole of the invagination, and the abutting ck twin-spot tissue in the top-left corner of the panel. (L) Malformation with a central depression and ftup2 macro and microchaetae. Red, sensillum campaniforme, as shown at higher magnification in the inset. In K and L, some of the mutant ftup bristles are coloured yellow.

View larger version (93K): [in this window] [in a new window]   Fig. 2. Expression of tup in the imaginal wing disc. (A) Early second instar disc. Green, Tup; red, aprK568-lacZ, a marker for the dorsal compartment. (B) Late second instar disc. (C) Notum region of a mid-third instar disc. Red, pnr-Gal4 UAS-lacZ. (D) Late third instar disc. Red, Ara/Caup. Dotted lines indicate position of the LN/WH and WH/WP borders. Asterisk, region of possible low accumulation of Tup. a, anterior; p, posterior; MN, medial notum; LN, lateral notum; PLN, posterior lateral-most notum; WH, wing hinge; WP, wing pouch; tg, tegula.

View larger version (90K): [in this window] [in a new window]   Fig. 3. Expression of wing-hinge markers in tup clones located in the notum region. tup clones are identified by the absence of GFP (green) expression. (A) First instar-induced tupisl-1 clone (asterisk). The lacZ insertion line l(2)09261 is derepressed. (B-B'') tup2 clones (arrowheads) derepressing zfh2 (B') and/or msh89-lacZ (msh, B''). Asterisk, endogenous msh expression in the notum. (C) Optical z-axis section through tupex4 clones. Red, msh89-lacZ; blue, Twist, a marker for adepithelial cells; arrowheads, nuclei of tupex4 cells; arrow, peripodial membrane. (D) Extruding tupex4 clone stained with phalloidin (red). Arrowhead, apical actin accumulation. (E) tup2 clone inducing msh-lacZ expression autonomously and non-autonomously (arrowhead). (F) First instar-induced tup2 clone with enhanced expression of msh-lacZ. Compare (arrowheads) with the wild-type disc (G). (H) Derepression of sal (purple) within a tupex4 clone (asterisk) and in cells surrounding it. Compare with wild-type sal expression (I).

View larger version (142K): [in this window] [in a new window]   Fig. 4. Effect of tup clones on notum genes. tup clones are identified by the absence of GFP (green) expression. (A) Wild-type expression of ush and pnr-Gal4 in the medial notum. Arrowheads, ush expression in the lateral notum and hinge regions. (B) First instarinduced tupex4 clone. pnr-Gal4 expression is lost. The Ush accumulation pattern (arrowhead) is similar to that in the hinge area. (C) Second instar-induced tupex4 clones. ush is repressed. (D) tup2 clones (blue arrowheads) repress pnr-Gal4 (blue or white) and two of the tup2 clones upregulate ara/caup (white arrowheads). (E) tup2 clones. Only that clone in the central notum (arrow) represses ara/caup (red). Proximal clones upregulate ara/caup (arrowheads). Inset shows that pnr-Gal4 expression (white or blue) persists in most clones. (F) Regions are outlined where tup clones lose (blue, mapped with 13 clones overlapping the area) or gain (white, mapped with 28 clones) ara/caup expression. (G) tupisl-1 clone. msh-lacZ is upregulated (arrowhead) and ara/caup downregulated (arrow) in the same cells. (H) Anterior tup2 clone. ara/caup (arrow) and msh-lacZ (arrowhead) are both upregulated in some cells of the clone. (I) First instar-induced tupex4 clone showing derepression of msh-lacZ and inhibition of eyg. Inset shows wild-type expression of eyg.

View larger version (67K): [in this window] [in a new window]   Fig. 5. Chip and Ssdp are required for notum specification. (A,B) Chipe5.5 clones (absence of green) lose Eyg (red in A), accumulate Zfh2 (blue), and non-autonomously upregulate msh (red in B, arrowheads). (C,D) Clones of either M+ Ssdpneo48 (C, absence of green) or UAS-Chip-expressing (D, green) cells lose Eyg (red) and accumulate Zfh2 (blue).

View larger version (103K): [in this window] [in a new window]   Fig. 6. Overexpression of tup and ara synergistically promote transformation towards notum. (A-D) Mesothoracic pleurae. Red arrowheads indicate anterior (anp) and posterior (pnp) notopleural, and ventral sternopleural (vsp) bristles; black arrowheads, tegulae (tg); asterisks, vertical clefts. (A) Wild type. (B) UAS-ara; C765-Gal4. (C) UAS-tup/C765-Gal4. Arrow, notum-like outgrowth on the vertical cleft. (D) UAS-ara; UAS-tup/C765-Gal4. Flies were grown at 17°C. Black arrow, notum expansion towards the pleura; wing is missing. White arrow, notum-like structure adjacent to sternopleural bristles (red arrowhead). (E-H) Wing discs showing expression of eyg (green) and DC-lacZ (red). Genotypes in E-H correspond with those shown in A-D, respectively. Arrowhead (G) indicates ectopic expression in the prospective pleura (this location was verified in an optical z-section). (H) eyg is expressed at the dorsal hinge (arrowhead) and the wing and pleura territories (arrows). Insets show red channel images of the pleura and wing pouch areas. (I) UAS-ara; UAS-tup/dppblk-Gal4 fly. Notum-like structures form on the central wing (asterisk), pleura (black arrow) and sternopleurite (red arrow). aw and pw, anterior and posterior parts of the wing, respectively. (J) UAS-ara; UAS-tup/dppblk-Gal4 mesothoracic leg disc. Arrowhead, ectopic eyg expression. (K-M) Wild-type (L) and UAS-ara; UAS-tup/dppblk-Gal4 (K,M) wing discs showing either eyg (K) or nub and zfh2 (L,M) expression. Arrows (K) indicate that eyg expression is expanded to the driver territory. Arrowheads (M) indicate that nub and zfh2 expression is lost from the driver territory.

View larger version (68K): [in this window] [in a new window]   Fig. 7. Regulation of tup in the wing disc. Red, Tup; blue or white, Ara/Caup. (A) M+ Egfr1K35 clones (absence of green) remove ara/caup expression (arrowheads) but do not inhibit tup (arrows). (B) M+ Ras85DC40b clones (absence of green) inhibit ara/caup (arrowheads), but not tup expression. (C) Second (top row) and early third (bottom row) instar discs. Overexpression of Mkp3 (green) inhibits ara/caup (arrowhead), but not tup (arrow). (D) Expression of ara/caup in wild-type discs of similar age to those shown in C. (E) Clones expressing UAS-Ras1V12 (green) activate ara/caup (arrowheads) in the wing hinge. tup is not activated, or only so at very low levels. (F) A tkva12 clone (absence of green) removes tup expression in the medial (arrowhead), but not in the lateral (arrow) notum. (G) Second (top) and early third (bottom) instar discs. Overexpression of UAS-Dad (green) blocks Tup accumulation (arrowheads). Compare with Fig. 2B,C. (H) Clones expressing UAS-tkvQD (green) activate tup in the medial (arrowhead), but not in the lateral (arrow) notum.

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