From: Subject: p53 and Pten control neural and glioma stem/progenitor cell renewal and differentiation : Article : Nature Date: Mon, 27 Oct 2008 08:37:54 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C9380F.4E7ED690" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C9380F.4E7ED690 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.nature.com/nature/journal/v455/n7216/full/nature07443.html p53 and Pten control = neural and glioma stem/progenitor cell renewal and differentiation : = Article : Nature

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Nature 455, 1129-1133 (23 October 2008) | = doi:10.1038/nature07443;=20 Received 2 May 2008; Accepted 10 September 2008

p53 and Pten control neural and glioma stem/progenitor cell = renewal=20 and differentiation

Hongwu Zheng1,12,=20 Haoqiang Ying1,12,=20 Haiyan Yan1,=20 Alec C. Kimmelman1,4,=20 David J. Hiller8,=20 An-Jou Chen1,=20 Samuel R. Perry1,2,=20 Giovanni Tonon1,=20 Gerald C. Chu1,2,5,=20 Zhihu Ding1,=20 Jayne M. Stommel1,=20 Katherine L. Dunn1,=20 Ruprecht Wiedemeyer1,=20 Mingjian J. You1,=20 Cameron Brennan9,10,=20 Y. Alan Wang1,2,=20 Keith L. Ligon1,3,5,6,=20 Wing H. Wong8,=20 Lynda Chin1,2,7=20 & Ronald A. DePinho1,2,11

  1. Department of Medical Oncology,
  2. Center for Applied Cancer Science, Belfer Foundation = Institute for=20 Innovative Cancer Science,
  3. Center for Molecular Oncologic Pathology, Dana-Farber = Cancer=20 Institute and Harvard Medical School, Boston, Massachusetts 02115, USA
  4. Harvard Radiation Oncology Program,
  5. Department of Pathology,
  6. Division of Neuropathology,
  7. Department of Dermatology, Brigham and Women's Hospital, = Harvard=20 Medical School, Boston, Massachusetts 02115, USA
  8. Department of Statistics, Stanford University, Stanford, = California=20 94305, USA
  9. Department of Neurosurgery, Memorial Sloan-Kettering = Cancer Center,=20 New York, New York 10065, USA
  10. Department of Neurosurgery, Weill-Cornell Medical = College, New=20 York, New York 10065, USA
  11. Departments of Medicine and Genetics, Harvard Medical = School,=20 Boston, Massachusetts 02115, USA
  12. These authors contributed equally to this work.

Correspondence to: Ronald A. DePinho1,2,11=20 Correspondence and requests for materials should be addressed to R.A.D.=20 (Email: ron_depinho@dfci.harvard.edu= ).

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Abstract

Glioblastoma (GBM) is a highly lethal brain tumour = presenting as=20 one of two subtypes with distinct clinical histories and molecular = profiles. The=20 primary GBM subtype presents acutely as a high-grade disease that = typically=20 harbours mutations in EGFR, PTEN and INK4A/ARF (also known = as=20 CDKN2A), and the secondary GBM subtype evolves from the slow = progression=20 of a low-grade disease that classically possesses PDGF and = TP53=20 events1,=20 2,=20 3.=20 Here we show that concomitant central nervous system (CNS)-specific = deletion of=20 p53 and Pten in the mouse CNS generates a penetrant = acute-onset=20 high-grade malignant glioma phenotype with notable clinical, = pathological and=20 molecular resemblance to primary GBM in humans. This genetic observation = prompted TP53 and PTEN mutational analysis in human = primary GBM,=20 demonstrating unexpectedly frequent inactivating mutations of = TP53 as=20 well as the expected PTEN mutations. Integrated transcriptomic = profiling,=20 in silico promoter analysis and functional studies of murine = neural stem=20 cells (NSCs) established that dual, but not singular, inactivation of = p53=20 and Pten promotes an undifferentiated state with high renewal = potential=20 and drives increased Myc protein levels and its associated signature. = Functional=20 studies validated increased Myc activity as a potent contributor to the = impaired=20 differentiation and enhanced renewal of NSCs doubly null for p53 = and=20 Pten (p53-/- Pten-/-) as well = as tumour=20 neurospheres (TNSs) derived from this model. Myc also serves to maintain = robust=20 tumorigenic potential of = p53-/- Pten-/- TNSs.=20 These murine modelling studies, together with confirmatory=20 transcriptomic/promoter studies in human primary GBM, validate a = pathogenetic=20 role of a common tumour suppressor mutation profile in human primary GBM = and=20 establish Myc as an important target for cooperative actions of p53 and = Pten in=20 the regulation of normal and malignant stem/progenitor cell = differentiation,=20 self-renewal and tumorigenic potential.

High-grade malignant glioma, the most common intrinsic = brain=20 tumour, is uniformly fatal despite maximum treatment3.=20 A wealth of molecular genetic studies has established central roles of = the=20 RTK-PI3K-PTEN, ARF-MDM2-p53 and INK4a-RB pathways in = gliomagenesis3,=20 4.=20 To explore the role of p53 and Pten in glioma, we used the = hGFAP-Cre=20 transgene5,=20 6=20 to delete p53 alone or in combination with Pten in all CNS = lineages using conditional p53 (ref. 7)=20 and Pten alleles (Supplementary=20 Figs 1 and 2a=96c). Because broad CNS deletion of Pten = results in=20 lethal hydrocephalus in early postnatal life (data not shown), modelling = efforts=20 henceforth emphasized the Ptenlox/+ genotype.

Clinically, between 15 to 40 weeks of ages, 42 out of 57 = (73%) of=20 the=20 hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 mice presented with acute-onset neurological symptoms=97seizure, ataxia = and/or=20 paralysis (Fig.=20 1a). Histopathologically, all 42 neurologically symptomatic mice = harboured=20 malignant gliomas that were classified on the basis of WHO (World Health = Organization) criteria8=20 as anaplastic astrocytomas (WHO III, n =3D 28, 66%) or GBM (WHO = IV,=20 n =3D 14, 34%; Fig.=20 1b). These GBMs had classical features of pseudopalisading necrosis, = marked=20 cellular pleomorphism, and highly infiltrative spread including = perineuronal and=20 perivascular satellitosis as well as subpial spread in the cerebral = cortex (Supplementary=20 Fig. 3a). Occasionally tumours had abnormal vessels suggestive of=20 microvascular proliferation. All tumours showed increased mitoses (Ki67=20 staining) and expression of the classical human glioma markers Gfap and = Nestin=20 (Fig.=20 1c). Necropsy of 15 neurologically asymptomatic mice showed no cases = of=20 incipient low-grade glioma disease but 8 had high-grade pathology = including very=20 small lesions with anaplastic features of nuclear atypia, multinucleated = tumour=20 cells and/or high cellularity (Supplementary=20 Fig. 3b). For the remaining genotypes, 4 out of 23=20 hGFAP-Cre+;p53lox/lox mice developed = anaplastic astrocytoma (WHO III); conversely 19 out of 23=20 hGFAP-Cre+;p53lox/lox mice, 12 out = of 12=20 hGFAP-Cre+;p53lox/+;Ptenlox/+ = mice and=20 10 out of 10 hGFAP-Cre+;p53lox/+ = mice had no=20 CNS pathology and developed only non-CNS tumours (data not shown).

Figure=20 1: p53 and Pten inactivation cooperate to induce=20 high-grade malignant gliomas.

a, Kaplan=96Meier tumour-free survival curves for mice of = indicated=20 genotypes as a function of weeks. '+' designates the wild-type allele,=20 'L' denotes the conditional allele. b, Graph shows = frequency and=20 grade of gliomas versus non-CNS malignancies observed in end-stage of = indicated=20 mice from a. Asy* indicates neurological asymptomatic=20 hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 mice (n =3D 15) killed for non-CNS malignancies. c, = Haematoxylin and=20 eosin (H&E) histology and immunohistochemical staining of sections = of WHO=20 grade III and IV malignant gliomas from=20 hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 mice with antibodies against Ki67, Gfap and Nestin. Scale bars, = 50 m.

<= A=20 class=3Dhigh-res=20 href=3D"http://www.nature.com/nature/journal/v455/n7216/fig_tab/nature074= 43_F1.html">High=20 resolution image and legend (306K)Download=20 Power Point slide (795K)

Slides may be downloaded for educational use, = according=20 to the terms described in Na= ture=20 Publishing Group's licensing policy.


Historically, TP53 inactivation has been = considered a=20 classical lesion in low-grade astrocytomas and secondary GBM but = infrequently in=20 primary GBM1,=20 9.=20 The remarkable clinical and histological resemblance of this model to = the=20 primary GBM subtype in humans prompted TP53 and PTEN = resequencing=20 in human primary GBM. Of the 35 clinically annotated human primary GBM = samples,=20 10 (29%) tumours registered prototypical TP53 mutations and 14 = (40%)=20 tumours had PTEN missense mutations, insertions, deletions or = splicing=20 mutations (Supplementary=20 Table 1). Moreover, six out of ten tumours with TP53 = mutations=20 harboured concomitant PTEN mutations or homozygous deletion.=20 Encouragingly, our mutational data agrees with The Cancer Genome Atlas = data=20 reporting TP53 and PTEN as the two most commonly mutated = tumour=20 suppressor genes (http://tcga-data.nci.nih.gov/= tcga/findArchives.htm).=20 These results, together with recent population-based studies10,=20 11,=20 indicate that TP53 is a key tumour suppressor for both GBM = subtypes.

Consistent with frequent PTEN loss of = heterozygosity (LOH;=20 60=9670%) in human high-grade glioma3,=20 16 out of 16 mouse high-grade gliomas showed no Pten expression in = tumour cells=20 but a robust signal in surrounding non-malignant cells and intratumoral = vessels=20 (Fig.=20 2a). Polymerase chain reaction (PCR) genotyping indicated that six = out of=20 seven tested tumours sustained loss of the wild-type Pten allele = (Fig.=20 2b). The Pten reduction to homozygosity and the documented=20 Cre-mediated deletion of both p53 floxed alleles indicate that = the=20 inactivation of both genes is required for gliomagenesis. Loss of Pten=20 expression correlated with activation of key PI3K signalling surrogates: = Akt and=20 ribosomal protein S6 kinase (Fig.=20 2c). In accordance with human high-grade disease, eight out of eight = malignant murine gliomas expressed high Vegf levels relative to normal = brain=20 tissue (Fig.=20 2c). Co-activation of multiple receptor tyrosine kinases in human = primary=20 GBM12=20 was also evident in the murine tumours with robust Pdgf=20 expression overlapping with strong regional activation of Egfr (Supplementary=20 Fig. 4a=96d).

Figure=20 2: hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 gliomas mirror key features of human malignant gliomas.

a, Pten expression is completely extinguished in tumour cells. = Sections of three independent malignant gliomas were stained with = haematoxylin=20 and eosin (H&E) and an anti-Pten antibody. 'N' indicates the = adjacent normal=20 regions of the tumour cells; the arrows point to Pten-positive vascular = cells=20 embedded in the tumour. b, The wild-type Pten allele is = lost in=20 glioma cells. Genomic DNA isolated from liver tissues and brain tumour = cells was=20 subjected to PCR-based assays for genotyping Pten and p53 = alleles.=20 '+' designates the Pten wild-type allele, 'L' denotes the=20 conditional allele, and 'D' denotes the inactivated form of the=20 conditional allele after Cre-mediated recombination. c,=20 Immunohistochemical staining of mouse normal brain or glioma sections = with=20 antibodies against activated phosphorylated Akt (pAkt), phosphorylated = ribosomal=20 protein S6 kinase (pS6) and Vegf. d, TNS lines isolated from = independent=20 malignant gliomas were cultured in NSC medium or differentiation medium = (1%=20 fetal bovine serum (FBS)) and immunostained for Nestin, Gfap and Tuj1 as = indicated. Scale bars, 50 m=20 (a, c); original magnification in d 400.

High=20 resolution image and legend (391K)Download=20 Power Point slide (775K)

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A classical feature of human high-grade malignant glioma = is a=20 significant degree of intertumoral and intratumoral morphological and = lineage=20 heterogeneity, hence the moniker glioblastoma 'mutliforme'. This = characteristic=20 plasticity was evident in the=20 hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 gliomas in which occasional tumours (5 out of 50) presented with both = astrocytic=20 and oligodendroglial histopathological features (Supplementary=20 Fig. 5). The basis for morphological variability is not known and = may=20 relate, among many possibilities, to the acquisition of an immature=20 developmental state with multipotency and/or differentiative plasticity. = Consistent with this notion, all murine tumours express stem or lineage=20 progenitor markers (including Nestin, Gfap and Olig2) similar to human = glioma=20 profiles13=20 but are negative for mature neuronal and oligodendrocyte markers (NeuN = and Mbp;=20 Supplementary=20 Fig. 6a, b). This stem/progenitor marker profile is in accord with = the=20 ability of all murine tumours tested to readily generate TNSs with (i) = strong=20 tumour-initiating potential with secondary tumours faithfully retaining = the=20 primary tumour's histological features (Supplementary=20 Fig. 7); (ii) robust NSC marker Nestin expression; and (iii) limited = capacity to differentiate into astrocytic and neuronal lineages after = exposure=20 to differentiation agents (Fig.=20 2d). As NSC/progenitor cells have been proposed to be the preferred=20 cell-of-origin for GBM6,=20 the immature marker profile and varied morphological presentation of our = murine=20 tumours prompted us to posit that Pten and p53 = deficiencies might=20 contribute to gliomagenesis by affecting NSC self-renewal and = differentiation=20 potential.

To explore this hypothesis, we characterized primary = murine=20 embryonic day (E)13.5 NSC cultures singly or doubly null for p53 = and=20 Pten. Compared to NSCs null for either Pten or p53, = which=20 show only modestly increased proliferation and self-renewal reflected by = neurosphere formation capacity14,=20 15,=20 16,=20 NSCs null for both p53 and Pten showed significant = proliferation=20 and self-renewal activity (Fig.=20 3a and Supplementary=20 Fig. 8a). This effect on NSC renewal, coupled with the = aforementioned varied=20 tumour histology, suggests that combined p53 and Pten loss = might=20 cooperate in tumorigenesis by impairing NSC differentiation potential. = When NSCs=20 were continuously cultured in NSC medium, all genotypes showed a similar = robust=20 expression of NSC/progenitor markers (for example Nestin) and minimal = expression=20 of differentiated lineage markers (Supplementary=20 Fig. 8b). After exposure to differentiation-inducing medium, = wild-type and=20 single-null NSC cultures differentiated into Gfap-positive astrocytes,=20 Tuj1-positive neurons, or O4-positive oligodendrocytes. In contrast,=20 p53-/- Pten-/- NSCs failed to respond = to these=20 differentiation cues and retained stem-cell-like morphology and lineage = marker=20 (Nestin) expression (Fig.=20 3b and Supplementary=20 Fig. 9a). Similar differentiation defect and abnormal self-renewal = potential=20 were also observed in adult NSCs that were deleted for Pten and=20 p53 postnatally (Supplementary=20 Fig. 9b, c). The contribution of Pten deficiency in = maintaining=20 impaired differentiation was further verified by the ability of the Akt=20 inhibitor triciribine17=20 to enable differentiation of NSCs null for both Pten and = p53 (Supplementary=20 Fig. 10a, b).

Figure=20 3: p53 and Pten coordinately regulate Myc protein = level as=20 well as NSC self-renewal and differentiation.

a, The number of neurospheres formed by=20 p53-/- Pten-/- NSCs in culture is=20 significantly increased as compared to wild-type or singly null NSCs; = asterisk,=20 P < 0.001; n =3D 3. Values represent = mean  s.d.=20 from three experiments. b, The multilineage differentiation = potential was=20 impaired in p53-/- Pten-/- NSCs. D, = DAPI=20 (blue); G, Gfap (green); N, Nestin (red); O4 (red); WF, white field; T, = Tuj-1=20 (red). c, Combined inactivation of p53 and Pten in = NSCs=20 stimulates Myc protein expression. d, Knockdown of Myc expression = restores p53-/- Pten-/- NSC = differentiation=20 capacity. Lower panel, western blot of double-null NSC Myc protein = expression=20 after infected with indicated lentiviral shRNA. Note Myc expression in=20 shMyc2- and shMyc3-infected=20 p53-/- Pten-/- cells is comparable to = that in=20 p53-/- cells, and shMyc1 as a control shows no=20 knockdown. Original magnification used for b and d: 200.

High=20 resolution image and legend (356K)Download=20 Power Point slide (738K)

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To understand the molecular basis of impaired = differentiation=20 capacity, we performed transcriptome comparisons of murine=20 p53-/- NSCs with=20 p53-/- Pten-/- NSCs at 1 day = after=20 exposure to the differentiation inducer. Among the 410 genes showing = significant=20 differential expression (Supplementary=20 Table 2) promoter analysis identified E2F and Myc motifs as two of = the most=20 enriched promoter binding elements (1.7=20 and 1.4,=20 respectively; P < 10-4). Notably, = promoter=20 analysis using 69 pretreatment human primary GBM cases in the TCGA = database=20 showed strong enrichment of MYC binding elements: 10=20 p53-/- Pten-/- double-mutant tumours = versus=20 the 59 remaining tumours (1.40,=20 P =3D 2.20  10-3)=20 or versus the 12 p53-/- single-mutant tumours = (1.46,=20 P =3D 1.54  10-3).

MYC is well-known for its roles in cell cycle = progression and=20 apoptosis18=20 as well as in stem cell self-renewal and differentiation during = development and=20 oncogenic processes19,=20 20,=20 21,=20 22.=20 It is also notable that both p53 and Pten/PI3K pathways can directly = regulate=20 MYC with p53 repressing MYC transcription by directly binding to = the=20 MYC promoter23,=20 whereas downstream PI3K pathway arms can modulate MYC translation and = protein=20 degradation24,=20 25.=20 In agreement, Myc protein levels were substantially increased in the = murine=20 double-null NSCs, but only marginally elevated in = p53-/- or=20 Pten-/- NSCs when compared to wild-type controls (Fig.=20 3c), raising the possibility that p53 and Pten cooperate to regulate = Myc=20 levels which in turn could control NSC self-renewal and = differentiation.

To test this hypothesis, we examined the effect of Myc = knockdown=20 on murine p53-/- Pten-/- NSC = differentiation=20 potential and observed that Myc short hairpin RNA (shRNA; shcMyc2 = and=20 shcMyc3), which reduced Myc levels to those in = p53-/-=20 NSCs, largely restored their differentiation capacity (Fig.=20 3d and Supplementary=20 Fig. 11a). Conversely, enforced Myc expression in=20 p53-/-NSCs repressed their differentiation and enabled = retention of stem/progenitor marker expression (Nestin and Sox2; Supplementary=20 Fig. 11b), indicating that the concomitant loss of p53 and Pten = elevates Myc=20 activity to impede NSC differentiation capacity.

The strong pleiotropic activities attributed to MYC = demands tight=20 control of its expression to avoid development of diverse human = malignancies,=20 including gliomas21,=20 26.=20 Our finding that the concomitant loss of p53 and Pten compromises NSC=20 differentiation capacity by means of elevated Myc levels prompted = further=20 assessment of its relevance in the so-called 'brain cancer stem cells' = in our=20 model. Using murine TNSs which are enriched for such tumour initiating = cells=20 (TICs), Akt inhibitor treatment strongly reduced Myc protein and = promoted=20 differentiation (Fig.=20 4a, b and Supplementary=20 Fig. 12a). Correspondingly, Myc knockdown in TNS cells not only = markedly=20 reduced their proliferation and self-renewal capacity (Fig.=20 4c and Supplementary=20 Fig. 12b, c) but also strongly sensitized them to differentiation = induction=20 (Fig.=20 4d and Supplementary=20 Fig. 12d). Notably, although ten out of ten intracranial injections = of=20 vector-transduced murine TNSs resulted in lethal infiltrating gliomas = within=20 1 month, nine out of ten mice injected with Myc knockdown TNSs = survived for=20 more than 3 months (Fig.=20 4e). Thus, Myc has a crucial involvement in maintaining the impaired = differentiation and potent tumorigenic potential of p53- and=20 Pten-inactive TNSs (Supplementary=20 Fig. 13).

Figure=20 4: Attenuated Myc expression restores=20 hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 TNS differentiation potential and reduces tumorigenic = potential.

a, Inhibition of the Akt pathway by triciribine induces TNS = cell=20 differentiation. Two independent TNS lines were cultured in 1% FBS = in the=20 absence or presence of triciribine (Tri, 5 M) for = 7 days before being subjected to immunostaining with antibodies = against=20 Nestin (Nes, red), Gfap (Green) and Tuj1 (red). D, DAPI (blue). = b,=20 Inhibition of the Akt pathway in TNS cells with triciribine attenuates = their=20 cellular Myc expression. c, Knockdown of Myc expression in TNS = cells=20 reduces their self-renewal potential assessed by sphere formation; = asterisk,=20 P < 0.001; n =3D 3. Values represent = mean  s.d.=20 from three experiments. d, shRNA-mediated reduction of Myc = expression in=20 TNS cells sensitizes cells to differentiation stimuli. Cells infected = with=20 control (shGfp) and the indicated shRNA were incubated with=20 differentiation medium before being subjected to lineage marker = analysis.=20 e, shRNA-mediated reduction of Myc expression represses TNS = tumorigenic=20 potency in orthotopically transplanted SCID mice. Original magnification = for=20 a and d: 200.

High=20 resolution image and legend (281K)Download=20 Power Point slide (666K)

Slides may be downloaded for educational use, = according=20 to the terms described in Na= ture=20 Publishing Group's licensing policy.


The identification of TICs with stem-like properties in = diverse=20 human cancers including GBM represents an important conceptual advance = in cancer=20 biology with therapeutic implications27,=20 28.=20 These TICs seem to constitute a reservoir of self-sustaining cells with = potent=20 tumorigenic potential. However, unlike normal NSCs which readily = differentiate=20 along a developmental hierarchy into lineage-restricted differentiated=20 progenies, TNSs derived from = p53-/- Pten-/-=20 malignant gliomas show resistance towards differentiation cues. The = diminished=20 tumorigenicity of these TICs on restoration of differentiation = potential, along=20 with recent reports supporting pro-differentiation as a potential = strategy to=20 inhibit GBM-derived TICs29,=20 30,=20 encourages the identification and testing of agents targeting these=20 differentiation pathways including MYC in the treatment of primary GBM = in=20 humans.

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Methods Summary

Standard gene targeting and chimaera formation methods = were used=20 to generate the conditional PtenL allele in the mouse = germ=20 line in which Pten exon 5 is flanked by loxP sites; the = conditional=20 p53L mouse was generated by A. Berns; hGFAP-Cre=20 mice were purchased from the Jackson Laboratory. All mice were = maintained in=20 pathogen-free facilities and followed for development of neurological = deficits.=20 After culling, tissues were collected and processed for histological,=20 immunohistochemical, immunofluorescence or western blot analyses, as = detailed in=20 the Methods. For p53 and Pten mutation analysis, = surgically=20 resected human primary glioblastoma were flash-frozen and genomic DNA = was=20 prepared from frozen tumour samples. For microarray analysis, total RNAs = from=20 indicated early passage NSCs were amplified and labelled by standard = methods and=20 hybridized to Affymetrix 430=20 2.0 chips. For tumorigenic analysis, TNS cells isolated from=20 hGFAP-Cre+;p53lox/lox;Ptenlox/+=20 murine malignant gliomas were infected with indicated lentivirus shRNA = and=20 orthotopically injected into the forebrain of SCID mice. Animals were = observed=20 daily for the development of neurological deficits and subjected to = histological=20 analysis once killed.

Full=20 methods accompany this paper.

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Supplementary Information

Supplementary=20 information accompanies this paper.

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Acknowledgements

We thank A. Berns for providing p53L = mice; S.=20 Zhou and S. Jiang for mouse husbandry and care; R. T. Bronson for = discussion on=20 pathology analysis; K. Montgomery for discussion on sequencing; and = Y.-H. Xiao,=20 B. Feng and J. Zhang for bioinformatic help. H.Z. was supported by Helen = Hay=20 Whitney Foundation. H. Ying is a recipient of the Marsha Mae Moeslein = Fellowship=20 from the American Brain Tumor Association. A.C.K. is a recipient of the = Leonard=20 B. Holman Research Pathway Fellowship. Z.D. is supported by the Damon = Runyon=20 Cancer Research Foundation. J.M.S. is supported by a Ruth L. Kirschstein = National Research Service Award Fellowship. R.W. is supported by a = Mildred=20 Scheel Fellowship (Deutsche Krebshilfe). Grant support comes from the = Goldhirsh=20 Foundation (R.A.D.), and NIH grants U01 CA84313 (R.A.D.), RO1CA99041 = (L.C.) and=20 5P01CA95616 (R.A.D., L.C., W.H.W., C.B. and K.L.L.). R.A.D. is an = American=20 Cancer Society Research Professor supported by the Robert A. and Renee = E. Belfer=20 Foundation Institute for Innovative Cancer Science.

Author Contributions H.Z. and H. Ying performed = the=20 experiments and contributed equally as first authors. R.A.D. supervised=20 experiments and contributed as senior author. M.J.Y. generated the=20 PtenL mouse allele. D.J.H., W.H.W. and G.T. conducted = the=20 microarray and promoter analyses. K.L.L., H.Z. and G.C.C. provided the = pathology=20 analyses. H. Yan, A.C.K., A.-J.C., S.R.P., Z.D., J.M.S, K.L.D. and R.W.=20 performed the experiments. C.B. contributed patient samples and = pathologic=20 information. L.C. and Y.A.W. contributed to the writing of the=20 manuscript.

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Online Methods

Mice

p53L and hGFAP-Cre mice = have been=20 described previously5,=20 7.=20 PtenL mice were generated using a standard knock-in = approach=20 in which Pten exon 5 is flanked by loxP sites (details on = targeting=20 construct and procedures are available on request). Mice were interbred = and=20 maintained on a FvB/C57Bl6 hybrid background in pathogen-free conditions = at the=20 Dana-Farber Cancer Institute, monitored for signs of ill-health every = other day,=20 and euthanized and necropsied when moribund. All manipulations were = performed=20 with Institutional Animal Care and Use Committee (IACUC) = approval.

Histology and immunohistochemistry

Once killed, mice were perfused with 4% = paraformaldehyde=20 (PFA) and brains were dissected, followed by overnight post-fixation in = 4% PFA=20 at 4 =B0C. Serial sections were prepared at 5 m for=20 paraffin sections or 10 m for=20 cryostat sections with every tenth slide stained by haematoxylin and = eosin=20 (DF/HCC Research Pathology Cores). Tumour grading was determined by = K.L.L. and=20 H.Z. on the basis of the WHO grading system for malignant = astrocytoma8.=20 Immunohistochemical and immunofluorescence analyses were performed as=20 described31.=20 The Pdgfr=20 and pEgfr double immunohistochemical staining was performed using DakoCytomation=20 EnVision doublestain system ( K1395,=20 Dako) following manufacturer's instructions. The primary antibodies used = were:=20 Ki67 ( VP-RM04,=20 Vector), Gfap ( Z0334,=20 DAKO), Gfap ( 556330,=20 BD Pharmingen), Nestin ( MAB353,=20 Chemicon; specifically for mouse), Nestin ( MAB5326,=20 Chemicon; specifically for human), Pten ( 9559,=20 Cell Signaling), phospho-AktSer473 ( 3787,=20 Cell Signaling), phospho-ribosomal protein S6 kinase ( 2215,=20 Cell Signaling), Cyclin D1 ( 18-0220,=20 ZYMED), Vegf ( sc-152,=20 Santa Cruz), Pdgfr (=20 3174,=20 Cell Signaling), phospho-PdgfrY754=20 ( sc-12911,=20 Santa Cruz), Egfr ( IHC-00005,=20 Bethyl), phospho-EgfrY1068 ( ab40815,=20 Abcam), phospho-EgfrY1173 ( sc-12351,=20 Santa Cruz), Olig-2 ( AB9610,=20 Chemicon), Tuj-1 ( MMS-435P,=20 Covance), O4 ( MAB1326,=20 R&D), NeuN ( MAB377,=20 Millipore), Mbp (ab7349, Abcam), Myc (ab39688, Abcam), and Cre ( 69050-3,=20 Novagen). Images were captured using a Leica DM1400B=20 microsystem and Leica FW4000=20 version 1.2.1.

Cell culture

Primary NSCs were isolated from the brain = subventricular=20 zone (SVZ) of E13.5 embryos or 1-month-old mice with the indicated = genotype as=20 previously described31,=20 32.=20 NSCs were maintained in NSC=20 proliferation media (05702, StemCell) supplemented with 20=20 ng ml-1 EGF=20 (E4127, Sigma) and 10 ng ml-1 bFGF=20 (F0291, Sigma). To generate primary TNS cells, tumour samples from = freshly=20 dissected mouse brains were subjected to mechanical and enzymatic = dissociation.=20 Single-cell suspensions were cultured in NSC proliferation media. Tumour = spheres=20 formed were then disaggregated and used for indicated assays. NSC=20 differentiation assays were carried out by plating the indicated cells = in=20 culture wells on coverslips precoated with 15 g = ml-1=20 poly-l-ornithine=20 (P3655, Sigma) and 1 g = ml-1=20 fibronectin=20 (F1141, Sigma); the cells were incubated in neurobasal medium = supplemented with=20 1% FBS for 7=9610 days, and the differentiation capacities were = examined=20 under either a light or fluorescence=20 microscope (Nikon). For TNS cell differentiation, cells were = incubated in=20 differentiation media with varying doses of triciribine=20 (BioMol) or vehicle ( dimethylsulphoxide,=20 Sigma). Knockdown of mouse Myc was performed by infecting the indicated = cells=20 with lentivirus containing shMyc constructs (provided by W. = Hahn). The=20 shRNA constructs shMyc1, shMyc2 and shMyc3 = correspond to=20 clone IDs TRCN000000 54856, 42517 and 42513, respectively (The = DFCI-Broad RNAi=20 Consortium, commercially available from Sigma-Aldrich).

Western, cell growth and self-renewal assays

Western blot assays were performed as previously=20 described31=20 with antibodies against Myc ( sc-42,=20 Santa Cruz), phospho-AktSer473 ( 9271,=20 Cell Signaling), Pten ( 9569,=20 Cell Signaling) and Actin ( sc-1615,=20 Santa Cruz). For in vitro cell growth assays, NSCs or TNS cells = (10,000)=20 were plated in triplicate in 96-well format and incubated in NSC = proliferation=20 media for 5 days, and growth was quantified using Luminescence=20 ATP detection assay system (PerkinElmer). Self-renewal capacity was = measured=20 by plating 1,000 cells per well (6-well plate) in NSC proliferation = media=20 containing EGF and bFGF with 0.3% agarose=20 (A9049, Sigma). The number of neurospheres or tumour neurospheres that = formed=20 subsequently per well was quantified after 10=9614 days and relative = sphere=20 formation was plotted versus indicated control. Three replicates were = performed=20 for each. All experiments were conducted at cell = passage <5.

Orthotopic transplants

Female=20 SCID mice (Charles River) aged 6=968 weeks were anesthetized = and placed=20 into stereotactic apparatus equipped with a Z axis (Stoelting). A = small=20 hole was bored in the skull 0.5 mm anterior and 3.0 mm lateral = to the=20 bregma using a dental drill. Twenty thousand cells in Hanks Buffered = Salt=20 Solution was injected into the right caudate nucleus 3 mm below the = surface=20 of the brain using a 10-l=20 Hamilton syringe with an unbeveled 30 gauge needle. The scalp was = closed=20 using a 9-mm Autoclip Applier. Animals were followed daily for the = development=20 of neurological deficits.

Mutation screening

Frozen tumour specimens were obtained from the = Memorial=20 Sloan Kettering Cancer Center tumour bank. Genomic DNA was prepared from = frozen=20 primary GBM tumour samples using the Qiagen genomic=20 purification kit. Coding exons were PCR amplified and sequenced = using=20 standard protocols at the Harvard Partners Center for Genetics and = Genomics as=20 previously described33.=20 All known single nucleotide polymorphisms and synonymous mutations were = removed=20 from the analysis in the current study. This study was approved by the=20 Institutional Review Board of the hospital.

Microarray analysis

Early passage wild-type and indicated mutant NSCs = were=20 incubated with NSC proliferation media or differentiation media for = 18 h.=20 RNA was isolated using Trizol=20 (Invitrogen) and the RNeasy=20 mini kit (Qiagen). Gene expression profiling was performed using the = Affymetrix 430=20 2.0 chips at DFCI Microarray core facility.

Promoter analysis

Gene expressions were modelled using dChip = software34.=20 Sets of genes differentially expressed pre- and post-differentiation = induction=20 were generated using the SAM statistic35,=20 with a cutoff of 2.0.=20 Promoter analysis on both these gene sets used the CisGenome software = (http://bi= ogibbs.stanford.edu/~jihk/CisGenome/index.htm)=20 to scan the 8 kb upstream to 2 kb downstream regions of these = genes=20 for the 550=20 motifs in the TRANSFAC 12.1 database. Enrichment was measured against = control=20 regions at a comparable distance from the transcription start sites of = random=20 genes.

Statistical analysis

Tumour-free survivals were analysed using Graphpad = Prism4.=20 Statistical analyses were performed using the non-parametric = Mann=96Whitney test.=20 Significance of enrichment in the promoter analysis was computed based = on=20 Poisson distribution with Bonferroni correction. Comparisons of cell = growth,=20 self-renewal and differentiation were performed using the unpaired = Student's=20 t-test. For all experiments with error bars, standard deviation = was=20 calculated to indicate the variation within each experiment and data, = and values=20 represent mean  s.d.

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ISSN: 0028-0836

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PADDING-LEFT: 15px; DISPLAY: inline; BACKGROUND: = url(/common/images/icons/next.gif) no-repeat 0px 0.3em; COLOR: #c03; = FONT-SIZE: 100%; TEXT-DECORATION: none } DIV.natjobmore A:visited { MARGIN: 0px; PADDING-LEFT: 15px; DISPLAY: inline; BACKGROUND: = url(/common/images/icons/next.gif) no-repeat 0px 0.3em; COLOR: #c03; = FONT-SIZE: 100%; TEXT-DECORATION: none } .box#natjob DIV.natjobmore A:hover { BACKGROUND-IMAGE: url(/common/images/icons/next_hover.gif); = TEXT-DECORATION: underline } .box#natjob DIV.natjobmore A:focus { BACKGROUND-IMAGE: url(/common/images/icons/next_hover.gif); = TEXT-DECORATION: underline } UL.natjob UL { PADDING-BOTTOM: 0px; LIST-STYLE-TYPE: none; MARGIN: 0px; PADDING-LEFT: = 0px; PADDING-RIGHT: 0px; FONT-SIZE: 100%; PADDING-TOP: 0px } P.natjobmore { PADDING-BOTTOM: 5px; MARGIN: 0px } P.nateventmore { PADDING-BOTTOM: 5px; MARGIN: 0px; PADDING-TOP: 5px } .box#natprod { BORDER-BOTTOM: #e7e7e7 1px solid; BORDER-LEFT: #e7e7e7 1px solid; = PADDING-BOTTOM: 6px; MARGIN: 10px 0px; PADDING-LEFT: 6px; PADDING-RIGHT: = 6px; BACKGROUND: #fff; COLOR: #000; BORDER-TOP: #e7e7e7 1px solid; = BORDER-RIGHT: #e7e7e7 1px solid; PADDING-TOP: 6px } H2.natprod { PADDING-BOTTOM: 5px; TEXT-TRANSFORM: lowercase; MARGIN: 0px 0px 0.75em; = PADDING-LEFT: 7px; PADDING-RIGHT: 7px; BACKGROUND: #d8d8d8; = LETTER-SPACING: -0.04em; FONT-SIZE: 120%; PADDING-TOP: 5px } H2.natprod A { TEXT-DECORATION: none } H2.natprod SPAN.first { COLOR: #f00 } H2.natprod SPAN.second { COLOR: #000 } .box#natprod FORM { MARGIN: 0px } DIV.natprod { BORDER-BOTTOM: black 1px solid; BORDER-LEFT: black 1px solid; = PADDING-BOTTOM: 4px; PADDING-LEFT: 4px; PADDING-RIGHT: 4px; BACKGROUND: = #f3f2f2; BORDER-RIGHT: black 1px solid; PADDING-TOP: 4px } DIV.natprodmore { PADDING-BOTTOM: 0.75em; PADDING-LEFT: 0px; PADDING-RIGHT: 0px; = PADDING-TOP: 0px } DIV.natprodmore A:link { PADDING-LEFT: 7px; DISPLAY: block; BACKGROUND: = url(/common/images/arrow_red.gif) no-repeat 0px 0.5em; COLOR: #c03; = 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