A Wingless and Notch double-repression mechanism = regulates G1[ndash]S transition in the Drosophila wing

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Subject Categories: Cell=20 Cycle | Develo= pment=20

The EMBO Journal advance online = publication 1=20 May 2008;
doi: 10.1038/emboj.2008.84

A Wingless and Notch double-repression = mechanism=20 regulates G1=96S transition in the Drosophila = wing

H=E9ctor Herranz¹, Lidia P=E9rez¹, = Francisco A Mart=EDn² and Marco Mil=E1n¹

¹ ICREA and Institute for = Research in=20 Biomedicine, Barcelona, Spain
² Centro de = Biolog=EDa Molecular CSIC-UAM, Madrid, Spain

To whom=20 correspondence should be = addressed
Marco Mil=E1n, ICREA=20 and Institute for Research in Biomedicine, Parc Cient=EDfic = de=20 Barcelona, Josep Samitier, 1-5, Barcelona 08028, Spain. = Tel.: +34 93=20 4034902; Fax: +34 93 4037109; E-mail: marco.milan@irbbarcelona.org =

Received 12=20 December 2007; Accepted 2 April 2008; Published online 1 May = 2008.=20

Abstract

The control of tissue growth and patterning is = orchestrated in various multicellular tissues by the = coordinated=20 activity of the signalling molecules Wnt/Wingless (Wg) and = Notch,=20 and mutations in these pathways can cause cancer. The role = of these=20 molecules in the control of cell proliferation and the = crosstalk=20 between their corresponding pathways remain poorly = understood.=20 Crosstalk between Notch and Wg has been proposed to organize = pattern=20 and growth in the Drosophila wing primordium. Here we = report=20 that Wg and Notch act in a surprisingly linear pathway to = control=20 G1=96S progression. We present evidence that these molecules = exert=20 their function by regulating the expression of the = dmyc=20 proto-oncogene and the bantam micro-RNA, which = positively=20 modulated the activity of the E2F transcription factor. Our = results=20 demonstrate that Notch acts in this cellular context as a = repressor=20 of cell-cycle progression and Wg has a permissive role in=20 alleviating Notch-mediated repression of G1=96S progression = in wing=20 cells.

Keywords: bantam, dMyc, E2F, = micro-RNA=20

Introduction

During development of multicellular = organisms,=20 Wnt/Wingless (Wg)- and Notch-signalling pathways are = involved in the=20 determination of a variety of cell fates and in the control = of=20 tissue growth (Bray,=20 2006; Clevers,=20 2006). In many cellular contexts, genetic manipulations = that=20 change the activities of these two proteins result in cancer = (Bienz=20 and Clevers, 2000; Radtke=20 and Raj, 2003). Thus, tight regulation of Notch and = Wnt/Wg is=20 crucial for proper development and survival of multicellular = organisms.

The wing primordium of Drosophila is = a very=20 suitable model system to define, at a genetic and cellular = level,=20 the role of these pathways in the development of highly=20 proliferative tissues. Cell-fate specification by Notch and = Wg is a=20 well-known and defined process; by contrast, little is known = about=20 their role in the control of cell proliferation. The wing = primordium=20 arises as a group of 30=9640 cells in the embryonic ectoderm = that=20 proliferates during the three larval stages to reach a final = size of=20 around 50 000 cells (Garc=EDa-Bellido=20 and Merriam, 1971; Madhavan=20 and Schneiderman, 1977). Early in development, the wing = becomes=20 subdivided into a dorsal (D) and a ventral (V) cell = population, or=20 compartment, by the activity of the LIM-Homedomain = transcription=20 factor Apterous in D cells (Diaz-Benjumea=20 and Cohen, 1993; Blair=20 et al, 1994), and cell interactions between D and = V cells=20 lead to activation of Notch at the compartment boundary (Diaz-Benjumea=20 and Cohen, 1995; de=20 Celis et al, 1996). During the third instar = larval stage,=20 Notch activity induces Wg expression at the dorsal=96ventral = (DV)=20 boundary (Diaz-Benjumea=20 and Cohen, 1995; de=20 Celis et al, 1996), and another set of cell = interactions=20 between boundary and nearby non-boundary cells takes place = to=20 maintain Notch activity and Wg expression at the DV boundary = (Figure=20 1). The DV boundary has a role in organizing growth of = the=20 whole-wing primordium (Diaz-Benjumea=20 and Cohen, 1993). Notch and Wg have been postulated as = being=20 responsible for this organizing activity, although data are=20 controversial (Giraldez=20 and Cohen, 2003; Johnston=20 and Sanders, 2003). Late in development, cells at the DV = boundary are characterized by cell-cycle arrest and define = the=20 so-called Zone of Non-proliferating Cells (ZNC; O'Brochta=20 and Bryant, 1985; Phillips=20 and Whittle, 1993). Wg has been reported to induce this=20 cell-cycle arrest (Johnston=20 and Edgar, 1998; Duman-Scheel=20 et al, 2004).

Figure=20 1
Wg- and Notch-mediated cell interactions = involved in=20 DV boundary formation. (A) Late third instar = wing=20 imaginal disc labelled to visualize Wg (green) and = Senseless=20 (red) protein expression in boundary and non-boundary = cells,=20 respectively. (B) Illustration describing the = cell=20 interactions that take place between boundary (white) = and=20 non-boundary (black) cells that lead to activation of = Notch=20 (N) and expression of Wg and Cut along the DV boundary = of the=20 Drosophila wing primordium. N, Notch; Dl, = Delta; Ser,=20 Serrate.

Here we have revised the role of Wg and = Notch in the=20 control of cell proliferation and present evidence that a Wg = and=20 Notch double-repression mechanism controls G1=96S transition = in the=20 wing primordium. Our results indicate that these signalling=20 molecules exert their function through regulation of the=20 bantam micro-RNA (Brennecke=20 et al, 2003) and the proto-oncogene dmyc = (Johnston=20 et al, 1999). Our work clarifies and simplifies = the role=20 of Notch and Wg in cell-cycle control in the = Drosophila wing=20 and provides a very suitable model by which to analyse the = function=20 of Notch- and Wg-signalling pathways in the regulation of = the=20 cell-cycle machinery.

Results and = discussion

The ZNC is defined by Notch = activity

Stable activation of Notch and expression of = Wg along=20 the DV boundary relies on a positive feedback loop between = boundary=20 and non-boundary cells (Figure=20 1). Boundary cells activate the receptor Notch and = express Wg,=20 whereas non-boundary cells respond to Wg and signal back by=20 expressing the Notch ligands Serrate and Delta. Boundary and = non-boundary cells are characterized late in development by = their=20 cell-cycle arrest and constitute the ZNC (O'Brochta=20 and Bryant, 1985). Anterior non-boundary cells are = arrested in=20 G2 by the activity of Wg (Johnston=20 and Edgar, 1998; Figure 8A). Wg exerts its function = through its=20 target genes achaete and scute, which inhibit = the=20 expression of cdc25/string, the universal eukaryotic=20 regulator of the G2/M transition. Posterior cells and = anterior=20 boundary cells are arrested in G1 (Johnston=20 and Edgar, 1998). This arrest has also been postulated = to be=20 defined by the activity of Wg (Johnston=20 and Edgar, 1998; Duman-Scheel=20 et al, 2004). Two observations suggest that = Notch, and=20 not Wg, is responsible for the cell-cycle arrest in G1. = First, the=20 Wg-signalling pathway is blocked in boundary cells by = activity of=20 Notch, as recently reported (Buceta=20 et al, 2007; Figure=20 1B). Second, some low-threshold target genes of Notch, = like=20 crumbs, are expressed in a broader domain that = includes both=20 boundary and non-boundary cells and corresponds to the ZNC = (Figure=20 2A and B; Herranz=20 et al, 2006). Here we have readdressed the role = of Wg and=20 Notch in this process.

Figure=20 2
The ZNC is defined by the activity of Notch. = Late=20 third instar wing discs labelled to visualize cells in = S-phase=20 by BrdU incorporation (antibody to BrdU in red or = white=20 (A, C, E, G, I, = N,=20 O, P)) or expressing the E2F1-responsive = reporter ORC1=96GFP (antibody to GFP in pink or = white=20 (B, D, F, H, J, = Q)).=20 (A, B) Wild-type wing discs labelled to visualize=20 crumbs=96lacZ expression (antibody to = -gal, green). Note failure to incorporate = BrdU (A) or=20 low levels of E2F1 activity (B) in those cells = expressing high=20 levels of the Notch-regulated gene crumbs, = which=20 corresponds to the ZNC. The DV compartment boundary is = marked=20 in this and other panels (d, dorsal; v, ventral). (C, = D)=20 Notch^ts2 wing discs reared at = the=20 restrictive temperature and labelled to visualize Wg = protein=20 expression (green). (E, F) ptc=96Gal4;=20 UAS=96Delta^DN (E) and=20 sal=96Gal4; = UAS=96mam^DN (F)=20 wing discs labelled to visualize Senseless (Sens, = green in E)=20 or Wg (green in F) protein expression. Sens expression = was=20 used to label the DV boundary. Red lines label the = domain of=20 Delta^DN or=20 mam^DN expression. (G, H) Wing = discs=20 with clones of cells lacking Su(H) activity=20 (Su(H)⁸), marked by the = absence of=20 GFP (G) or -gal (H) in green. The ZNC is marked by = horizontal=20 lines in G. (I) Wing disc with clones of cells lacking = arrow activity = (arrow²) marked=20 by the absence of GFP. The ZNC is marked by horizontal = lines.=20 (J) dpp=96Gal4 Gal80ts;=20 UAS=96TCF^DN wing disc = reared at=20 29=B0C during 30 h and labelled to visualize Wg = (green) protein=20 expression. The dpp=96Gal4 domain is marked by = arrowheads.=20 (K=96R) Wild-type (K, N), = C96=96Gal4;=20 UAS=96TCF^DN (L, O),=20 C96=96Gal4; UAS = =96TCF^DN=20 UAS=96N^intra (M, P, Q) = and=20 C96=96Gal4; UAS=96GFP (R) wing = discs labelled=20 to visualize Cut (green, K=96M) or GFP (green, R) = protein=20 expression, BrdU incorporation (red, N=96P) or = activity of the=20 E2F1 responsive reporter ORC1=96GFP = (antibody to=20 GFP in pink, Q). The ZNC is marked by horizontal = dashed lines.=20

The ZNC is characterized by its failure to = incorporate=20 bromodeoxyuridine (BrdU) (O'Brochta=20 and Bryant, 1985) or by the reduced activity levels of = the E2F1=20 transcription factor (Johnston=20 and Edgar, 1998; Figure=20 2A and B). E2F proteins, such as Drosophila E2F1, = regulate the expression of a number of genes required for = S-phase,=20 and their activity is inhibited by the Retinoblastoma (Rb) = proteins=20 like Drosophila Rbf (Rb-familiy protein, reviewed by = Dyson,=20 1998). The Rb pathway is a key regulator of the G1=96S = transition=20 and Drosophila Rbf is required for G1 arrest in the = ZNC (Duman-Scheel=20 et al, 2004). Activity of E2F1, labelled as dE2F = in the=20 Figures, can be visualized by observing the expression of = the=20 E2F1-responsive reporter ORC1=96GFP (Asano=20 and Wharton, 1999). It consists of the ORC1 = promoter=20 bearing functional E2F-binding sites driving the expression = of an=20 unstable Ftz=96GFP=96Myc fusion protein. As the half-life of = this=20 protein is short, this reporter monitors newly synthesized = protein=20 and permits examination of cell-cycle-regulated = transcription in=20 vivo.

We first analysed the role of Notch in = defining the=20 ZNC. In a temperature-sensitive Notch = loss-of-function=20 background (N^ts2) reared for 48 h = at=20 restrictive (29=B0C) temperature, cells at the ZNC = incorporated BrdU=20 and expressed ORC1=96GFP (Figure=20 2C and D). Blocking Notch activation by expression of a=20 dominant-negative form of the Notch ligand Delta or the = nuclear=20 Notch effector Mastermind (Giraldez=20 et al, 2002) caused similar effects (Figure=20 2E and F). Clones of cells located at the ZNC and mutant = for the=20 Suppressor of Hairless (Su(H)) transcription factor, the = nuclear=20 mediator of Notch signalling (Bray,=20 2006), incorporated BrdU and expressed the = E2F1-responsive=20 reporter (Figure=20 2G and H; see also Supplementary Figure S1). We used a=20 fluorescence-associated cell sorter (FACS) to collect data = on the=20 cellular DNA content of dissociated, GFP-sorted, ZNC wing = disc=20 cells, and to confirm the role of Notch in defining the G1 = block.=20 Blocking Notch signalling in these cells led to a decrease = in the=20 fraction of cells in G1 (Supplementary Figure S1). Altogether, = these=20 results imply that the activity of the Notch-signalling = pathway is=20 required for the cell-cycle arrest in G1 that takes place at = the=20 ZNC.

Wg has been reported to be required for the = G1 arrest=20 at the ZNC, as blocking Wg signalling in all wing margin = cells=20 causes cells to enter into S-phase (in C96;=20 UAS=96TCF^DN larvae; Figure=20 2N and O; and Johnston=20 and Edgar, 1998; Duman-Scheel=20 et al, 2004; see also Figure=20 2R for the pattern of expression of the C96=96Gal4 = driver). We=20 further analysed the requirement of the Wg pathway in this = process.=20 Unexpectedly, clones of cells mutant for arrow, the = Wg=20 co-receptor (Wehrli=20 et al, 2000), and located at the ZNC, did not = incorporate=20 BrdU (Figure=20 2I; Supplementary Figure S1; 74 out of 84 = clones=20 located at the ZNC did not incorporate BrdU; n = (scored=20 discs)=3D33). Similarly, expression in a subset of ZNC cells = of a=20 dominant-negative form of the LEF-1/TCF transcription factor = (TCF^DN), the nuclear mediator of canonical Wg = signalling=20 (Logan=20 and Nusse, 2004), did not increase BrdU nor E2F1 = activity levels=20 either (Figure=20 2J; Supplementary Figure S1). Altogether, = these=20 results indicate that Wg signalling is not required for the=20 cell-cycle arrest in G1 that occurs at the ZNC. The effects = reported=20 on G1 progression after blocking Wg signalling in all = wing-margin=20 cells (Johnston=20 and Edgar, 1998; Duman-Scheel=20 et al, 2004) can be explained by the Notch- and=20 Wg-dependent positive feedback loop that operates at the DV = boundary=20 (Figure=20 1B), which may also compromise the activity of Notch in = these=20 conditions. Indeed, expression of the Notch-regulated genes=20 cut and wg was reduced (Figure=20 2K and L; and data not shown), and increasing Notch = activity=20 levels, by means of expression of an activated form of the = Notch=20 receptor (N^INTRA), counteracted the = effects=20 of TCF^DN expression and restored cut = expression=20 (Figure=20 2M), reduced E2F activity (Figure=20 2Q) and BrdU incorporation (Figure=20 2P). Blocking Wg signalling only in a subset of = wing-margin=20 cells did not compromise Notch activity, as reported by = expression=20 of cut and wingless (Figures=20 3H, 4F=20 and F'). This might be caused by the fact that the = Notch- and=20 Wg-dependent positive feedback loop that operates at the DV = boundary=20 is not affected in these conditions. Note also that = TCF^DN=20 expression in these experiments was temporally controlled by = the=20 Gal4/Gal80ts system (see Materials and methods for details) = and=20 induced during the third instar stage (30 h before = dissection) to=20 circumvent the requirement of Wg signalling in this feedback = loop.

Figure=20 3
Regulation of E2F activity by Notch and Wg.=20 (A=96H) = Abruptex^M1 (A,=20 B), = dpp=96Gal4;UAS=96N^intra=20 (C, C', D, D'), ap=96Gal4 = Gal80ts,=20 UAS=96GFP; = UAS=96TCF^DN wing=20 (E, F), ap=96Gal4 Gal80ts, UAS=96GFP; = UAS=96Axin (G) and dpp=96Gal4 = Gal80ts;=20 UAS=96TCF^DN (H) late = third instar=20 wing discs labelled to visualize cells in S-phase by = BrdU=20 incorporation (antibody to BrdU in red or white (A, C, = C', E,=20 G)) or expressing the E2F1-responsive reporter=20 OCT1=96GFP (antibody to GFP in pink or = white (B,=20 D, D', F, H)). crb=96lacZ (antibody to = -Gal in green (A, B)), Wingless (Wg, green = (C, C', D,=20 D', H)) and GFP (green (E, G)) protein expression is = shown.=20 Panels C' and D' show higher magnifications of the = discs in=20 panels C and D. The anterior=96posterior (ap) boundary = is=20 labelled to show the cell-autonomous block in G1 = imposed by=20 the activity of N. N^intra = activity is=20 labelled by the expression of Wg (red line) and the G1 = block=20 by reduced levels of BrdU and dE2F activity. = (I,=20 J, K) ap=96Gal4, Gal80ts,=20 UAS=96GFP; = UAS=96TCF^DN (I),=20 wild-type (J) and ap=96Gal4 Gal80ts,=20 UAS=96GFP; = UAS=96N^intra (K)=20 early third instar wing discs labelled to visualize = BrdU=20 incorporation (red or white) and GFP (green) = expression. The=20 DV compartment boundary is marked in all panels (d, = dorsal; v,=20 ventral). Wing discs shown in panels E=96I were reared = at 29=B0C=20 during 24 h.

Figure=20 4
A Wg and Notch double-repression mechanism = regulates=20 G1 progression. (A=96D) = dpp=96Gal4,=20 UAS=96GFP;=20 = UAS=96N^intraUAS=96E2F=20 (A), ap=96Gal4, Gal80ts, UAS=96GFP;=20 UAS=96TCF^DN, UAS=96E2F = (B),=20 ptc=96Gal4, Gal80ts, UAS=96GFP;=20 UAS=96N^intra, UAS=96CycE = (C) and=20 ap=96Gal4, Gal80ts, UAS=96GFP;=20 UAS=96TCF^DN, UAS=96CycE = (D) late=20 third instar wing discs labelled to visualize cells in = S-phase=20 by BrdU incorporation (antibody to BrdU in red or = white) and=20 GFP (green) protein expression. (E=96G)=20 ap=96Gal4, Gal80ts, UAS=96GFP; = UAS=96TCF^DN=20 (E) and dpp=96Gal4, Gal80ts, UAS=96GFP;=20 UAS=96TCF^DN (F, F', G) = late third=20 instar wing discs labelled to visualize Wingless = protein (Wg,=20 red or white, E, G), wg mRNA (F), = wg=96lacZ=20 expression (antibody to -gal, F') and GFP (green, E) protein = expression.=20 Arrowheads in panels F, F' and G indicate the = dpp=96gal4=20 expression stripe. (H) ap=96Gal4, Gal80ts, = UAS=96GFP;=20 UAS=96TCF^DN,=20 UAS=96mam^DN late third instar = wing disc=20 labelled to visualize cells in S-phase by BrdU = incorporation=20 (antibody to BrdU in red or white) and GFP (green) = protein=20 expression. (I) Wing disc with clones of cells = lacking=20 arrow activity (arrow²), marked by the = absence of=20 GFP in green, and labelled to visualize Wg expression = (red or=20 white). Note ectopic expression of Wg in the clones = (red=20 arrows). (J) ap=96Gal4, Gal80ts,=20 UAS=96GFP;=20 = UAS=96TCF^DN,UAS=96mam^D=
N=20 late third instar wing discs labelled to visualize = cells=20 expressing the E2F1-responsive reporter = OCT1=96GFP=20 (antibody to GFP in pink or white) and Wg (green) = protein=20 expression. Wing discs shown in panels B and D=96J = were reared=20 at 29=B0C during 30 h. The DV compartment boundary is = marked in=20 all panels (d, dorsal; v, ventral). =

A G1=96S checkpoint controlled by a Wg = and Notch=20 double-repression mechanism

Notch activity is required in ZNC cells to = induce=20 cell-cycle arrest in G1, most probably by a reduction in = E2F1=20 activity levels (Duman-Scheel=20 et al, 2004). We then examined whether ectopic = activation=20 of Notch could reduce E2F1 activity levels and cause = cell-cycle=20 arrest in the rest of the wing cells, the so-called = wing-blade cells=20 (Figure=20 1A). For this purpose, we monitored these parameters in = wing=20 discs mutant for the Notch gain-of-function allele=20 Abruptex^M1 or expressing an = activated form=20 of the Notch receptor (N^INTRA). In = both=20 cases, the ZNC was expanded (Figure=20 3A=96D), as revealed by the reduced activity levels of = E2F1 and=20 the failure to incorporate BrdU. Notch induced this = cell-cycle=20 arrest in all cells located in the presumptive wing blade = and at any=20 time during the third instar larval stage (Figure=20 3C and K). Reduction in the activity of E2F1 mediated = this=20 cell-cycle arrest, as overexpression of E2F (known to bypass = the=20 negative effects of Rbf protein) restored cell-cycle = progression, as=20 monitored by BrdU incorporation (Figure=20 4A). To confirm that these cells were arrested in G1, we = coexpressed N^INTRA together with = cyclin E,=20 the key regulator and rate-limiting factor controlling = G1=96S=20 transition in wing cells (Knoblich=20 et al, 1994; Neufeld=20 et al, 1998). Interestingly, these cells = incorporated=20 BrdU (Figure=20 4C, compare with Supplementary Figure S2). Coexpression = of=20 cdc25/string, the rate-limiting factor of G2=96M = transition,=20 was not able to rescue the failure to incorporate BrdU (Supplementary Figure S2). Altogether, = these=20 results indicate that high levels of Notch induce a G1 block = by=20 reducing the activity of E2F1. We noted that high levels of = Notch=20 activity were able to induce an increase in BrdU = incorporation and=20 E2F activity in the proximal part of the wing (wing hinge, = data not=20 shown), and this increase might be the cause of the = overgrowth=20 phenotype observed in Abruptex^M1 = wing=20 discs.

Notch activation along the DV boundary = induces Wg=20 expression. Wg protein is secreted and forms a long-range = protein=20 gradient that reaches every wing-blade cell (Zecca=20 et al, 1996; Neumann=20 and Cohen, 1997; Strigini=20 and Cohen, 2000). We then analysed the role of Wg in = G1=96S=20 progression in these cells. For this purpose, we blocked the = Wg=20 pathway by expressing the dominant-negative form of TCF=20 (TCF^DN) or axin, and monitored BrdU incorporation = and=20 E2F1 activity levels. TCF^DN or axin expression = was=20 temporally controlled by the Gal4/Gal80ts system (see = Materials and=20 methods for details) and induced during the third instar = stage (30 h=20 before dissection) to circumvent the requirement of Wg = signalling in=20 cell survival and wing-fate specification (Giraldez=20 and Cohen, 2003; Johnston=20 and Sanders, 2003). We chose to use strong Gal4 drivers = to=20 completely block the Wg pathway. Interestingly, high levels = of=20 expression of TCF^DN or axin caused cell-cycle = arrest in=20 wing-blade cells. Those cells unable to transduce the Wg = signal did=20 not incorporate BrdU (Figure=20 3E and G; Supplementary Figure S1), and in these = cells the=20 expression levels of the E2F1-responsive reporter = ORC1=96GFP=20 were reduced (Figure=20 3F and H). All cells located in the presumptive wing = blade and=20 at any time during the third instar larval stage required = activation=20 of the Wg pathway to pass through S-phase (Figure=20 3E, G and I; compare with Figures=20 2A and 3J).=20 This cell-cycle arrest was due to a reduction in E2F1 = activity=20 levels, as coexpression of E2F rescued the cell = proliferation=20 defects (Figure=20 4B). Expression of cyclin E, and not = cdc25/stg,=20 was able to restore cell-cycle progression in the presence = of=20 TCF^DN (Figure=20 4D; Supplementary Figure S2). Taken = together, these=20 results indicate that Wg signalling is required in wing = blade cells=20 to reach the appropriate levels of E2F1 activity to trigger = G1=20 progression.

We noticed that reduced Wg signalling was = also able to=20 compromise the mitotic activity of wing blade cells (Supplementary Figure S2). We then = analysed the=20 possible regulation of the G2=96M transition in these = conditions. In=20 wing cells expressing TCF^DN, coexpression of=20 cdc25/string, the rate-limiting factor of G2=96M = transition,=20 was not able to rescue mitotic activity and BrdU = incorporation (Supplementary Figure S2). In contrast, = expression of cyclin E was able to rescue both of = them (Supplementary Figure S2; Figure=20 4D). Thus, the cell-cycle arrest imposed by lack of Wg=20 signalling takes place only in G1.

Notch- and Wg-signalling pathways exert = opposite=20 effects on G1=96S transition. Notch imposes G1 arrest = whereas Wg is=20 required for G1=96S progression. Interestingly, Notch is = known to be=20 repressed by the activity of Wg at different levels of its = pathway=20 during wing development (Axelrod=20 et al, 1996; Rulifson=20 et al, 1996; Micchelli=20 et al, 1997; de=20 Celis and Bray, 1997; Neumann=20 and Cohen, 1998). Consistent with this, blocking the Wg = pathway=20 in the dorsal compartment or in a stripe along the=20 anterior=96posterior compartment boundary induced ectopic = Notch=20 activation, as monitored by the expression of Wg protein, = wg=20 mRNA and wg=96lacZ (Figure=20 4E=96G), and clones of cells mutant for arrow, = the Wg=20 co-receptor, and located far away from the DV boundary = express Wg=20 (Figure=20 4I). These observations suggest that the opposite = effects of=20 Notch and Wg in the G1=96S transition could be explained by = a=20 Wg-mediated repression of the Notch pathway and a consequent = alleviation of the G1 block. To test this, we blocked the = Notch=20 pathway in cells unable to transduce the Wg signal and = analysed=20 their capacity to progress through G1. Interestingly, wing = cells=20 expressing the dominant-negative form of the nuclear Notch = effector=20 Mastermind (Mam^DN), together with = TCF^DN in the=20 dorsal compartment, incorporated BrdU and increased E2F1 = activity=20 levels (Figure=20 4H and J, compare with Figure=20 3E and F). Note that the expression pattern of Wg is = restored=20 (compare Figure=20 4E and J). These results imply that a Wg and Notch=20 double-repression mechanism controls G1=96S transition in = wing=20 cells.

dMyc and bantam micro-RNA mediate = regulation of=20 G1=96S by Wg and Notch

The proto-oncogene dMyc promotes = G1=96S=20 progression in Drosophila cells (Johnston=20 et al, 1999) and the cell-cycle arrest in G1 that = occurs=20 at the ZNC is defined by the absence of dMyc = expression (Johnston=20 et al, 1999; Duman-Scheel=20 et al, 2004; Figure=20 5A and B). We then analysed the regulation of = dMyc=20 expression by the activities of Notch and Wg. Interestingly, = and=20 consistent with the above results, downregulation of = dMyc=20 expression at the ZNC is defined by the activity of Notch, = and not=20 Wg (Johnston=20 et al, 1999; Duman-Scheel=20 et al, 2004). Blocking Notch activation by a=20 dominant-negative form of Mastermind (Giraldez=20 et al, 2002), or by using a=20 Notch^ts2 loss-of-function = background raised=20 at the restrictive temperature during 48 h, led to = expression of=20 dMyc at the ZNC (Figure=20 5C=96E). The effects reported on dMyc expression = after=20 blocking Wg signalling in all ZNC cells (Johnston=20 et al, 1999; Duman-Scheel=20 et al, 2004; see also Figure=20 5F) can be explained again by compromised Notch = activity, due to=20 the Wg and Notch positive feedback loop that operates at the = DV=20 boundary. Increasing Notch activity levels was able to = counteract=20 the effects of TCF^DN expression (Figure=20 5G).

Figure=20 5
Regulation of dMyc expression by Notch and = Wg.=20 (A=96N, P) Wild type (A, = B),=20 Notch^ts2 reared at the = restrictive=20 temperature for 48 h (C), ap=96Gal4, = UAS=96GFP;=20 UAS=96mam^DN (D),=20 C96=96Gal4; = UAS=96mam^DN (E,=20 H), C96=96Gal4; = UAS=96TCF^DN=20 (F), C96=96Gal4;=20 = UAS=96TCF^DNUAS=96N^intr=
a=20 (G), C96=96Gal4;=20 UAS=96mam^DN=20 UAS=96dMyc^dsRNA (I),=20 dpp=96Gal4; = UAS=96N^intra=20 (J, K), ap=96Gal4, Gal80ts, = UAS=96GFP;=20 UAS=96TCF^DN (L, N)=20 dpp=96Gal4; UAS=96Wg (M), and=20 ap=96Gal4, Gal80ts, UAS=96GFP;=20 UAS=96TCF^DN,=20 UAS=96mam^DN (P) late = third=20 instar wing discs labelled to visualize dMyc protein = (blue or=20 white (A, D, E=96J, M, N, P)) or dMyc mRNA (B, = C, K, L)=20 expression, and Wingless (Wg, red (A, E, J, M)), Cut = (red (F,=20 G)), Distalless (Dll, red (N, P)) or GFP (green (D, N, = P))=20 protein expression. The DV compartment boundary in = panels A=20 and B is indicated by a dashed line. Note dMyc=20 expression in GFP-expressing cells abutting the DV = compartment=20 boundary in D (compare with A) as well as in ZNC cells = lacking=20 Notch activity (C, E, H). (O, Q, = R)=20 ap=96Gal4, Gal80ts, UAS=96GFP;=20 UAS=96TCF^DN, UAS=96dMyc = (O),=20 dpp=96Gal4; UAS=96N^intra, = UAS=96dMyc=20 (Q, R) late third instar wing discs labelled to = visualize=20 cells in S-phase by BrdU incorporation (antibody to = BrdU in=20 red or white (O, Q)), or expressing the = E2F1-responsive=20 reporter OCT1=96GFP (pink (R)). GFP (O) and dMyc (R) = protein=20 expression (green) is also shown. Arrowheads in panels = Q and R=20 indicate the dpp=96gal4 expression stripe. Wing = discs=20 shown in panels L and N=96P were reared at 29=B0C = during 30 h.=20 (S) Wild-type wing disc labelled to visualize = dMyc=20 (blue) and Distalless (Dll, red) protein expression.=20

Expression of activated forms of the = receptor Notch=20 (N^INTRA), or the transcription = factor=20 Suppressor of Hairless (Su(H)-VP16), led to a reduction in=20 dMyc expression levels in wing-blade cells (Figure=20 5J and K; and Supplementary Figure S3). Ectopic = expression of=20 Wg in these cells did not cause such a reduction (Figure=20 5M). The cell-cycle arrest in G1 that takes place in = cells=20 expressing N^INTRA was at least in part due to a = reduction=20 in dMyc expression levels, as coexpression of dMyc induced = cells to=20 enter into the S-phase, as shown by BrdU incorporation (Figure=20 5Q). This observation correlates with the upregulation = of E2F1=20 activity levels in these cells (Figure=20 5R). We then examined the contribution of reduced levels = of dMyc=20 in the G1 block that occurs at the ZNC. To address this, we = analysed=20 the ability of reduced dMyc expression to impose a G1 block = at the=20 ZNC in a situation of reduced Notch activity. In wing discs=20 expressing a dominant-negative form of Mastermind along the = DV=20 boundary, dMyc expression and BrdU incorporation were = increased in=20 ZNC cells (Figure=20 5H). Decreasing dMyc protein levels in these cells, by = means of=20 a dMyc RNA interference construct, restored the ZNC, as = revealed by=20 the absence of BrdU incorporation (Figure=20 5I). These results indicate that reduced dMyc levels = caused by=20 the activity of Notch contribute to define the G1 block at = the=20 ZNC.

As reported above, a Wg and Notch = double-repression=20 mechanism induces G1=96S transition in wing cells not = located at the=20 ZNC. This mechanism correlates with the presence of = dMyc=20 expression, as wing cells unable to transduce the Wg signal=20 (expressing TCF^DN) showed reduced levels of = dMyc=20 (Figure=20 5L and N) and coexpression of Mam^DN, together = with=20 TCF^DN-rescued dMyc expression levels (Figure=20 5P). More interestingly, regulation of G1=96S transition = by this=20 double-repression mechanism was mediated by the presence of = dMyc, as=20 coexpression of dMyc together with TCF^DN rescued = G1=96S=20 progression, as visualized by BrdU incorporation (Figure=20 5O). We noticed that the Wg and Notch double-repression=20 mechanism did not regulate the expression of other = Wg-regulated=20 genes like Distalless, as wing cells unable to = transduce the=20 Wg signal lost expression of Distalless independently = of=20 Mam^DN coexpression (Figure=20 5N and P, compare with Figure=20 5S).

Altogether, these results indicate that dMyc = mediates=20 the activities of Notch and Wg in the regulation of G1=96S = transition=20 in the wing disc, defining the G1 cell-cycle arrest at the = ZNC and=20 facilitating G1 progression in the rest of cells. = Interestingly,=20 there was a striking correlation between dMyc = expression and=20 the activity of the bantam micro-RNA, which controls=20 cell-cycle progression in the Drosophila wing (Figures=20 6A and 7A;=20 Brennecke=20 et al, 2003). For this reason, we then analysed = the role=20 of bantam in this process. To monitor bantam = activity,=20 we used a green fluorescent protein (GFP) bantam = sensor that=20 expresses GFP under the control of a ubiquitously active = tubulin=20 promoter and has two perfect bantam target sites in = its 3'=20 untranslated region (Brennecke=20 et al, 2003). When present, the bantam = micro-RNA=20 reduces GFP expression through its RNA interference (RNAi) = effect.=20 The expression pattern of GFP is thus a negative image of = the=20 activity pattern of bantam micro-RNA. Brennecke=20 et al (2003) showed that the ZNC is characterized = by=20 higher levels of the bantam sensor (Figure=20 6A), and ectopic expression of bantam induces ZNC = cells=20 to enter into S-phase, as shown by BrdU incorporation. We = also=20 monitored E2F activity levels in these cells and verified = that=20 levels of E2F activity rose upon bantam expression in = ZNC=20 cells (Figure=20 6M), thereby indicating that absence of bantam=20 contributes to inducing G1 cell-cycle arrest through = regulation of=20 E2F1 activity.

Figure=20 6
Regulation of bantam activity by Notch = and Wg.=20 (A=96G; I, J) Wild = type (A,=20 A', E), ap=96Gal4;=20 UAS=96mam^DN (B),=20 dpp=96Gal4; = UAS=96N^intra=20 (C, C'), dpp=96Gal4; UAS=96Wg = (D),=20 C96=96Gal4; = UAS=96TCF^DN=20 (F), C96=96Gal4;=20 = UAS=96TCF^DNUAS=96N^INTR=
A=20 (G), dpp=96Gal4;=20 UAS=96TCF^DN (I) and=20 dpp=96Gal4; = UAS=96TCF^DN,=20 UAS=96mam^DN (J) late = third instar=20 wing discs labelled to visualize expression of the=20 bantam sensor (blue and white), and BrdU = incorporation=20 (red (A)), Cut (red (A')), Gal4 (red (B)), Wg (red (C, = C', I))=20 or Sens (red (D=96G, J)) protein expression. Note = symmetric=20 expression of the bantam sensor in dorsal and = ventral=20 cells abutting the DV compartment boundary (dashed = line) in=20 panels A and A', and lack of expression of the = bantam=20 sensor in dorsal cells abutting the DV compartment = boundary=20 (dashed line) in panel B (see also magnification). = Note=20 elevated levels of expression of the bantam = sensor in=20 the stripe of dpp=96Gal4 expression (red = arrowheads) in panels C and I, when compared with = panels A, B,=20 D and J. (H, K=96M) = C96=96Gal4;=20 = UAS=96TCF^DN/bantam¹ (H), dpp=96Gal4;=20 UAS=96TCF^DN,=20 UAS=96ban=96GFP (K), dpp=96Gal4;=20 UAS=96N^intra, = UAS=96ban=96GFP (L) and=20 dpp=96Gal4; UAS=96ban (M) late third instar = wing discs=20 labelled to visualize BrdU incorporation (red and = white H, K,=20 L) or E2F activity (red or white, M). Cut (green, H), = GFP=20 (green, K, L), Sens (blue, K) and Wg (blue, L, M) = protein=20 expression is also shown. Wing discs shown in panels = I=96K were=20 reared at 29=B0C during 30 h. =

Figure=20 7
Crosstalk between dMyc and bantam. (A) = Wild-type wing disc labelled to visualize the = bantam=20 sensor (blue) and dMyc protein expression (red and = white).=20 (B) ptc=96Gal4; = UAS=96bantamGFP wing=20 disc showing dMyc expression (red and white) and GFP = (green).=20 (C, D) Clones of cells lacking = bantam=20 activity (ban¹) marked by the absence of = GFP. dMyc=20 protein expression is shown in red or white. Note dMyc = expression in the clones (red arrows). = (E=96G)=20 sal=96Gal4; UAS=96dMyc (E),=20 dpp=96Gal4;=20 UAS=96N^intra,=20 UAS=96dMyc (F), and sal=96Gal4;=20 UAS=96dMyc^dsRNA (G) wing = discs=20 labelled to visualize expression of the bantam = sensor=20 (blue) and dMyc protein (red in E, red or white in G). = The DV=20 compartment boundary is marked in panels E and F (d, = dorsal;=20 v, ventral). (H) ptc=96Gal4;=20 UAS=96dMyc^dsRNA,=20 UAS=96bantamGFP late third instar wing = disc=20 labelled to visualize BrdU incorporation (red), GFP = (green)=20 and dMyc (blue or white) protein expression. = (I)=20 C96=96Gal4; UAS=96bantam late = third instar=20 wing disc, grown at 18=B0C and labelled to visualize = expression=20 of the bantam sensor (blue), BrdU incorporation = (red)=20 and dMyc protein expression (white).=20

We then analysed the regulation of = bantam=20 activity by Notch and Wg. Blocking Notch activation by means = of a=20 dominant-negative form of Mastermind (Giraldez=20 et al, 2002) reduced the expression of the = bantam=20 sensor at the ZNC (Figure=20 6B), and expression of an activated form of the Notch = receptor=20 (N^INTRA) led to an increase in its=20 expression levels in wing-blade cells (Figure=20 6C and C'). The activity of Notch is not mediated by Wg, = as Wg=20 ectopic expression did not cause reduction of bantam = activity=20 levels (Figure=20 6D), and Wg signalling was not required for repression = of=20 bantam activity in ZNC cells, either. Again, the = reported=20 effects of Wg signalling on bantam activity (Brennecke=20 et al, 2003) can be explained by compromised = Notch=20 activity, as increased Notch activity levels were able to = counteract=20 the effects of TCF^DN expression on bantam = activity=20 levels (Figure=20 6E=96G).The cell-cycle arrest in G1 that occurs in cells = expressing N^INTRA was at least, in part, due to a = reduction in bantam expression levels, as = coexpression of=20 bantam induced cells to enter into the S-phase, as = shown by=20 BrdU incorporation (Figure=20 6L). We then examined the contribution of reduced = bantam=20 activity in defining the G1 block that takes place at the = ZNC. In=20 wing discs with reduced Notch signalling along the DV = boundary, BrdU=20 incorporation was increased in ZNC cells (Figure=20 5H). We noted that Notch activity levels were not = completely=20 reduced in these conditions, as Wg and Cut were still = expressed in=20 some cells at the DV boundary (Figures=20 5E and 6H).=20 Thus, this background might be able to respond to changes in = the=20 levels of bantam activity. Indeed, halving the dose = of=20 bantam restored the ZNC, as revealed by the absence = of BrdU=20 incorporation (Figure=20 6H). These results indicate that reduced bantam = activity=20 contributes to define the G1 block at the ZNC.

In the rest of the wing cells, a Wg and = Notch=20 double-repression mechanism controls G1=96S transition. = There is again=20 a striking correlation between the activity levels of = bantam=20 and this double-repression mechanism. Indeed, wing cells = unable to=20 transduce the Wg signal (expressing TCF^DN) showed = increased expression of the bantam sensor (Figure=20 6I), and coexpression of Mam^DN, together with = TCF^DN, restored the expression levels of the=20 bantam sensor back to the endogenous ones (Figure=20 6J). More interestingly, regulation of the G1=96S = transition by=20 this double-repression mechanism was mediated by the = presence of=20 bantam, as coexpression of bantam, together = with=20 TCF^DN, rescued G1 cell-cycle progression, as = visualized=20 by BrdU incorporation (Figure=20 6K).

dMyc and bantam micro-RNA are two = independent=20 effectors of Notch and Wg

Notch imposed a G1 arrest at the ZNC by = reducing the=20 activity of bantam and the expression of dMyc. = Wg=20 repressed Notch and facilitated G1 progression in the rest = of wing=20 cells through dMyc and bantam. Expression of either = of these=20 genes was sufficient to induce G1 progression and bypass the = effects=20 of Notch or absence of Wg. We then questioned whether these=20 effectors are independently regulated by Notch and Wg, or,=20 alternatively, whether their expression depends on each = other.

Overexpression of bantam induced an = increase in=20 dMyc protein expression in wing-blade cells and at the ZNC = (Figure=20 7B, compare with Figure=20 7A), and overexpression of dMyc caused an increase in = the=20 activity levels of bantam (Figure=20 7E). By contrast, clones of cells mutant for = bantam did=20 not show reduction in dMyc protein levels (Figure=20 7C and D), and reduced levels of dmyc did not=20 considerably affect bantam activity levels (Figure=20 7G). Altogether, these results indicate that dMyc and=20 bantam are able, when overexpressed, to modulate each = other's=20 expression or activity levels, but are independently = regulated by=20 Notch and Wg.

Are dMyc and bantam two independent = effectors=20 regulating G1 progression? The evidence for dMyc is clear, = as in the=20 presence of Notch or absence of Wg signalling, dMyc = expression had=20 the capacity to rescue G1 progression (Figure=20 5M, O and P), and this happened in the presence of = reduced=20 levels of bantam activity. Note that dMyc = overexpression was=20 able to increase bantam activity levels only very = mildly at=20 the ZNC where Notch activity is high (Figure=20 7E), and was completely unable to restore the low = activity=20 levels of bantam imposed by Notch activation (compare = Figures=20 6C and 7F).=20 By contrast, bantam might rescue G1 progression = through=20 regulation of dMyc (Figure=20 7B). Note that in this case bantam is able to = bypass the=20 effect of Notch at the ZNC and induce an increase in dMyc = expression=20 (Figure=20 7B). We then analysed the capacity of bantam to = induce G1=20 progression after reducing dMyc expression levels. = Expression of=20 bantam, together with a dmyc double-stranded = RNA=20 (dsRNA) construct, induced G1=96S progression in wing cells = (Figure=20 7H), and this happened in the presence of very low = levels of=20 dMyc (Figure=20 7H). Altogether, these results imply that bantam = and dMyc=20 are two independent effectors of Wg and Notch that regulate = G1=20 progression.

These effectors are able to substitute each = other when=20 overexpressed in situations of reduced Wg signalling or high = levels=20 of Notch activity. However, their endogenous levels do not = appear to=20 do so, as loss of dMyc or bantam activity in = clones of=20 cells compromises cell-cycle progression (Johnston=20 et al, 1999; Brennecke=20 et al, 2003), and reduced levels of either = dMyc or=20 bantam, were able to define the ZNC (Figures=20 5 and 6).=20 This suggests that the endogenous levels of both effectors = are=20 rate-limiting. Interestingly, coexpression of both effectors = at=20 endogenous, wing-blade levels was able to drive G1=96S = progression in=20 ZNC cells. As shown in Figure=20 7I, ZNC cells with wing-blade levels of bantam = activity=20 and dMyc protein were able to enter S-phase, as monitored by = BrdU=20 incorporation. Thus, wing cells require the activity of both = bantam and dMyc to drive G1=96S = transition.

Concluding remarks

Here we have analysed the role of Notch and = Wg in the=20 regulation of G1=96S progression in the Drosophila = wing and=20 have provided evidence that a surprisingly linear pathway, = based on=20 a Wg and Notch double-repression mechanism, controls this = transition=20 (Figure=20 8B). Notch activity autonomously blocks cells in G1 and = defines=20 the ZNC at the DV boundary of the Drosophila wing. Wg = activity is required in wing-blade cells to repress Notch = signalling=20 and allow cells to enter the S-phase. These signalling = molecules=20 exert their function in a linear pathway through the = bantam=20 micro-RNA and the dmyc proto-oncogene, two = independent=20 effectors controlling the activity of the E2F1 transcription = factor=20 and regulate G1-to-S progression. Notch is known to be = repressed by=20 the activity of Wg at different levels of its pathway (Axelrod=20 et al, 1996; Rulifson=20 et al, 1996; Micchelli=20 et al, 1997; de=20 Celis and Bray, 1997; Neumann=20 and Cohen, 1998), but the level at which Wg represses = Notch=20 activity to facilitate dMyc expression, bantam = activity and G1=96S progression remains to be = elucidated.

Figure=20 8
Different effects of Notch and Wg in = cell-cycle=20 control in the Drosophila wing. (A) = Illustration=20 describing the control of the G2-to-M transition in = the=20 anterior (a) compartment by the activity of Wg through = its=20 target genes achaete and scute=20 (ac=96sc). ac=96sc are = known to=20 repress the expression of Drosophila cdc25 = string=20 (stg) in non-boundary cells. Note Wg does not = exert any=20 function in this process in posterior (p) cells. Notch = represses ac=96sc expression in boundary = cerlls,=20 this facilitating G2 progression in these cells. = (B)=20 Illustration describing the control of the G1=96S = transition by=20 the activities of Notch (N) and Wingless (Wg) at the = ZNC and=20 in wing-blade cells, through the downstream effectors=20 bantam and dMyc. =

There is evidence in mammals that Notch can = act as a=20 tumour-suppressor gene or as an oncogene depending on the = cellular=20 context (reviewed by Radtke=20 and Raj, 2003). The cellular context should then = modulate the=20 differential response of the cells to changes in Notch = activity, in=20 some cells leading to hyper-proliferation, and in others to=20 quiescence. How this context and the differential response = are=20 defined at the molecular level remains unclear. = Drosophila,=20 again, provides a very suitable model system to address this = issue.=20 Although our results highlight that Notch is involved in the = inhibition of G1=96S progression in wing discs, in other = developmental=20 contexts Notch exerts a proliferative function (Baonza=20 and Freeman, 2005; Firth=20 and Baker, 2005). Interestingly, in both cases, Notch = exerts an=20 effect through Rbf and E2F to positively or negatively = control the=20 G1=96S transition. We, therefore, speculate that the = molecular context=20 is then defined by the effectors available, like dMyc and=20 bantam, or by the presence of nuclear factors that = act as a=20 switch in the ultimate activation/inactivation of E2F = through=20 Rbf.

It is interesting to note that there are = three=20 consensus sequences for binding of Su(H) to DNA in the = dMyc=20 gene (data not shown), and in the particular case of human = T-cell=20 acute lymphoblastic leukaemia/lymphoma, c-myc = proto-oncogene=20 acts as a direct downstream target of Notch1 (Sharma=20 et al, 2006; Weng=20 et al, 2006). These results raise the possibility = that=20 dMyc might be a direct target of Notch in the=20 Drosophila wing. However, the fact that dMyc = is=20 negatively regulated by Notch and an activated form of Su(H) = (Su(H)-VP16) is able to repress dMyc expression, = suggests=20 that Notch regulates dMyc through a Notch target gene = that=20 may act as a repressor.

The DV boundary behaves as an organizer of = growth in=20 the developing wing, as ectopic DV boundaries induce = non-autonomous=20 proliferation of the surrounding tissue, and loss of the DV = boundary=20 leads to absence of wing tissue (Diaz-Benjumea=20 and Cohen, 1993). Notch has been shown to mediate this=20 organizing activity (Irvine=20 and Wieschaus, 1994; de=20 Celis et al, 1996), and previous reports showed = that Wg=20 is not sufficient to promote growth in the absence of an = endogenous=20 DV boundary (Klein=20 and Arias, 1998; Baena-Lopez=20 and Garcia-Bellido, 2003; Zecca=20 and Struhl, 2007). These results are consistent with the = data=20 obtained in this work and suggest that the organizing = activity of=20 the DV boundary should not be attributed to the activity of = Wg. Our=20 results indicate that the organizing activity of Notch is = strictly=20 non-autonomous, as Notch induces cell-cycle arrest in a=20 cell-autonomous manner. In this context, Wg signalling = alleviates=20 Notch-mediated repression of G1=96S progression in wing = cells. The=20 role of Wg can be defined as permissive rather than = instructive, as=20 wing cells unable to respond to Notch and Wg do proliferate. = Altogether, these results indicate that unknown signalling = molecules=20 regulated by Notch might exert the organizing activity of = the wing=20 DV boundary.

Materials and = methods

Drosophila strains

crb^{M11. M2} = (crb=96lacZ=20 in the text), UAS=96Ser^DN and=20 UAS=96Dl^DN (Herranz=20 et al, 2006); E2F1-responsive reporter (Asano=20 and Wharton, 1999), UAS=96dE2F1 UAS=96DP = (UAS=96E2F in=20 the text; Neufeld=20 et al, 1998, a gift from B Edgar), = UAS=96dMyc (Johnston=20 et al, 1999, a gift from B Edgar), = ban¹, ban=96sensor=96GFP and=20 UAS=96bantam=96GFP (Brennecke=20 et al, 2003, a gift from S Cohen), and=20 UAS=96dMyc^RNAi (Vienna Drosophila = RNAi=20 Center), UAS=96mam^DN (Helms=20 et al, 1999). Other stocks are described in = Flybase.

Antibodies

Guinea pig anti-Senseless (Nolo=20 et al, 2000, a gift from H Bellen), mouse anti-Wg = and=20 mouse anti-BrdU are described in the Developmental Studies = Hybridoma=20 Bank; rabbit anti--gal (Cappel) was also used. Anti-dMyc antibody = was raised=20 in guinea pig against recombinant full-length dMyc protein. = The=20 specificity of the anti-dMyc antibody was tested in wing = discs=20 overexpressing dMyc and in wing discs in which dMyc = expression is=20 reduced by means of a dsRNA construct of dmyc (see Supplementary Figure S4). BrdU = staining,=20 antisense digoxigenin-labelled RNA probes of dMyc and = wg mRNAs and in situ hybridization was = performed as=20 described by Milan=20 et al (1996).

Spatiotemporal gene expression = targeting in=20 Drosophila

We made use of the TARGET system developed = by McGuire=20 et al (2004). Adult flies carrying the Gal4 = drivers=20 ptc=96Gal4, dpp=96Gal4 or ap=96Gal4 and = the=20 thermosensitive version of the Gal4 repressor Gal80 = (Gal80ts) under=20 the control of the tubulin promoter = (tubulin=96Gal80ts)=20 were crossed with transgenic flies bearing one or more UAS=20 transgenes. Flies were allowed to lay eggs over a period of = 24 h at=20 18=B0C. The progeny was then grown at 18=B0C to maintain the = Gal4/UAS=20 system in a switched-off condition and transferred to 29=B0C = for a=20 range of periods during larval development to induce=20 Gal4/UAS-dependent expression.

Larval genotypes

The TARGET system:

Supplementary data

Supplementary data are available at The = EMBO=20 Journal Online (http://www.embojournal.org/).

Acknowledgements

We thank H Bellen, S Campuzano, S Cohen, B = Edgar, C=20 Estella, M Llimargas, E Mart=ED, G Morata, J-P Vincent, R = Wharton, the=20 Bloomington Stock Center and the Developmental Studies = Hybridoma=20 Bank for flies and reagents; L Johnston, Laura Buttitta, BA = Edgar=20 and three anonymous reviewers for comments on the = manuscript; and T=20 Yates for help with manuscript preparation. HH was funded by = a Juan=20 de la Cierva postdoctoral contract (Ministerio de = Educaci=F3n y=20 Ciencia), and MM laboratory was funded by a grant from the = Direcci=F3n=20 General de Investigaci=F3n Cient=EDfica y T=E9cnica = (BFU2004-00167/BMC), a=20 Grant from the Generalitat de Catalunya (2005 SGR 00118) and = by=20 intramural funds.

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	Abstract

	Introduction

	Results=20 and discussion

	Materials=20 and methods

	Acknowledgements

	References

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