From: Subject: Inhibitor of differentiation 4 drives brain tumor-initiating cell genesis through cyclin E and notch signaling -- Jeon et al. 22 (15): 2028 -- Genes and Development Date: Sat, 16 Aug 2008 16:42:25 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0006_01C8FFBF.0FFB6830" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0006_01C8FFBF.0FFB6830 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://genesdev.cshlp.org/cgi/content/full/22/15/2028 Inhibitor of differentiation 4 drives brain = tumor-initiating cell genesis through cyclin E and notch signaling -- = Jeon et al. 22 (15): 2028 -- Genes and Development
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RESEARCH = COMMUNICATION=20

Inhibitor of differentiation 4 drives brain tumor-initiating cell = genesis=20 through cyclin E and notch signaling

Hye-Min=20 Jeon1, Xun Jin1, = Joong-Seob=20 Lee1, Se-Yeong Oh1, = Young-Woo=20 Sohn1, Hyo-Jung Park1, = Kyeung=20 Min Joo2, Woong-Yang Park3,=20 Do-Hyun Nam2, Ronald A.=20 DePinho4, Lynda Chin4, and=20 Hyunggee Kim1,5= =20

1 School of Life Sciences and Biotechnology, Korea = University,=20 Seoul 136-713, Republic of Korea; 2 Department of = Neurosurgery,=20 Sungkyunkwan University School of Medicine, Seoul 135-710, Republic of = Korea;=20 3 Department of Biochemistry and Molecular Biology, Seoul = National=20 University School of Medicine, Seoul 110-799, Republic of Korea; = 4=20 Department of Medical Oncology and Center for Applied Cancer Science, = Belfer=20 Institute for Innovative Cancer Science, Dana-Farber Cancer Institute, = Harvard=20 Medical School, Boston, Massachusetts 02115, USA=20


3D"   =20 Abstract
Top
3D"Abstract
3D"Results and Discussion
3D"Materials and methods
Acknowledgments
3D"References =
 
Cellular=20 origins and genetic factors governing the genesis and = maintenance of=20 glioblastomas (GBM) are not well understood. Here, we report = a=20 pathogenetic role of the developmental regulator Id4 = (inhibitor of=20 differentiation 4) in GBM. In primary murine = Ink4a/Arf=96/=96=20 astrocytes, and human glioma cells, we provide evidence that = enforced=20 Id4 can drive malignant transformation by stimulating = increased=20 cyclin E to produce a hyperproliferative profile and by = increased=20 Jagged1 expression with Notch1 activation to drive astrocytes = into a=20 neural stem-like cell state. Thus, Id4 plays an integral role = in the=20 transformation of astrocytes via its combined actions on = two-key cell=20 cycle and differentiation regulatory molecules.

[Keywords: Id4; Ink4a/Arf=96/=96 astrocyte; notch = signaling;=20 cyclin E; neural stem-like cells; glioblastoma]

Received March 3, 2008; revised version accepted June 16, 2008.

The self-renewal, differentiative, and proliferative states = of tissue=20 stem and progenitor cells are maintained by a variety of = well-defined=20 cell fate determinant factors (Keller 20053DGo). Disregulation of such key developmental = regulators=20 and acquisition and maintenance of an immature = differentiation=20 phenotype are common themes across a broad spectrum of cancer = types,=20 including primary brain cancers (Garraway and Sellers 20063DGo). Indeed, a growing body of evidence supports = the view=20 that developmental regulators play direct roles in driving = aspiring=20 cancer cells toward a malignant phenotype, one that is = endowed with=20 stem-like cell properties including robust renewal potential = (Beachy=20 et al. 20043DGo). This cellular phenotype, embodied in the cancer = stem=20 cell paradigm (Pardal et al. 20033DGo), has been well documented in the highly = malignant=20 brain cancer, primary glioblastoma multiforme (GBM) (Galli et = al.=20 2004; Singh et al. 20043DGo). The cellular origins and specific genetic = elements=20 involved in the genesis and maintenance of these so-called = brain=20 cancer stem cells (BCSCs) remain areas of intensive=20 investigation.


3D"   =20 Results and Discussion
Top
Abstract
3D"Results and Discussion
3D"Materials and methods
Acknowledgments
3D"References=20
 

Id4 is = overexpressed=20 human GBM, and can drive transformation of murine = Ink4a/Arf=96/=96=20 astrocytes

Given the biological significance of stem cell fate determinants = in the regulation of normal self-renewal and differentiation, = we hypothesized that the genetic or epigenetic alterations in = classical cell fate determinants might cooperate with = prototypical=20 oncogenes and tumor suppressor genes to both effect malignant = transformation and endow stem cell-like renewal activity in=20 these tumors (Leung et al. 20043DGo;=20 Ligon et al. 20073DGo). We were particularly focused on the = expression and=20 functional activity of inhibitor of differentiation 4 (Id4) = in GBM=20 given its prominent role in governing neural stem cell (NSC) = fate=20 decisions (Yun et al. 20043DGo).

Id4 mRNA levels were found to be increased in nine of 15 GBM=20 samples as compared with human normal brain tissue = (Supplemental=20 Fig. S1A) and abundant Id4 mRNA and protein levels were = detected=20 in five of seven human glioma cell lines (A172, LN229, LN18,=20 U87MG, and T98G) and an immortalized human NSC line (HB1.F3)=20 (Cho et al. 20023DGo), but not in the normal human astrocytes (NHA)=20 (Supplemental Fig. S1B). Tumor microarray analysis showed 38 = of=20 68 newly diagnosed human GBM (WHO Grade IV) specimens with = robust Id4=20 expression with heterogenous staining patterns within the = same tumor=20 sample (Supplemental Fig. S1C). This heterogeneity and = importance of=20 Id family members in endothelial cell biology (Lyden et al. = 19993DGo)=20 prompted coimmunofluorescence studies to evaluate expression = of Id4=20 and the endothelial cell marker, CD31. The majority of=20 Id4+ cells were CD31=96 and, notably, = these=20 Id4+ cells were found to reside near the vasculature=97a = location postulated as the preferred residence of BCSCs = (Calabrese=20 et al. 20073DGo). In contrast, only a few Id4+ cells were=20 CD31+, and most of these positive cells were = associated=20 with macrovessels in the human GBM specimens (Supplemental = Fig.=20 S2).

These expression findings raised the possibility that Id4 = overexpression=20 might be relevant to the genesis and maintenance of a large=20 fraction of GBM cases, prompting us to determine whether = enforced=20 Id4 expression could drive malignant transformation of = primary=20 murine astrocytes derived from Ink4a/Arf=96/=96,=20 p53=96/=96, or wild-type mice. Ectopic Id4 = expression in=20 Ink4a/Arf=96/=96, but not wild-type or = p53=96/=96,=20 astrocytes significantly accelerated cell proliferation in = cell=20 culture (Supplemental Fig. S1D; data not shown) and enabled = tumor=20 formation in vivo (n =3D 6 mice) (Supplemental Fig. = S1E). Thus,=20 Id4 can function as a potent oncogene in astrocytes deficient = for the=20 signature GBM tumor suppressor, Ink4a/Arf.


Id4 drives conversion of Ink4a/Arf=96/=96 = astrocytes into=20 neural stem-like cells

Embryonic expression patterns of Id4 differ significantly from = the=20 other three Id family members (Id1, Id2, and Id3) in that Id4 = expression is primarily restricted to the developing central = nervous=20 system (Jen et al. 19963DGo). Disruption of the Id4 gene in mice results = in neural=20 tube proliferation defects, premature neuroblast = differentiation, and=20 overall reduction in brain size with drastic disruption in = brain=20 architecture (Bedford et al. 20053DGo;=20 Yun et al. 20043DGo). Accordingly, we observed that Id4 mRNA = levels are=20 high in the developing cerebral cortex, remaining relatively = constant=20 from embryonic day 13.5 (E13.5) to E19.5, and then declining = markedly=20 in postnatal life (Supplemental Fig. S3A). This Id4 = expression=20 profile is consistent with its role as a critical regulator = of=20 NSC/progenitor cell biology and cerebral cortex = development.=20

The defining properties of BCSCs include cancer-initiating = ability=20 upon implantation, extensive self-renewal potential, = sequential=20 neurosphere formation capacity, impaired multilineage = differentiation=20 potential, and capacity to generate nontumorigenic progeny = (Vescovi=20 et al. 20063DGo). To assess the potential of Id4 to commandeer the=20 mature differentiation state of astrocytes, we audited = astrocyte=20 (GFAP and S100β) and NSC (Nestin, Cd133, and Sox2) = markers=20 following transduction of Ink4a/Arf=96/=96 = astrocytes=20 with vectors encoding Id4-EGFP or EGFP only (designated = hereafter=20 as Id4-transduced or vector-transduced cells). As shown in Figure = 1A and Supplemental Figure S3, B and C, the expression = levels=20 of GFAP and S100β declined markedly, while all NSC = markers=20 increased dramatically in Id4-transduced = Ink4a/Arf=96/=96=20 astrocytes relative to vector-transduced control cells. = Furthermore,=20 upon placement of low numbers of Id4- or vertor-transduced=20 Ink4a/Arf=96/=96 astrocytes (one cell per square = millimeter at=20 the 24-well culture plates) into NSC culture media, the = former showed=20 increased capacity to produce neurosphere-like aggregates (Fig. = 1B).=20 Specifically, 43 neurospheres were generated in 38 of 96=20 single-cell-seeded wells in the Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes compared with seven = neurospheres=20 in 96 single-cell-seeded wells for the control cells (data = not=20 shown). Id4-transduced neurospheres possessed = Nestin+ and=20 Cd133+ cells (Fig. = 1C) and=20 exhibited self-renewal activity as single dissociated cells, = i.e.,=20 these cells could be serially passaged under the NSC culture=20 conditions (Fig.=20 1D). Thus, Id4 appears capable of driving=20 Ink4a/Arf=96/=96 astrocytes toward a = dedifferentiated=20 phenotype with characteristic NSC/progenitor markers and = renewal=20 activity.



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  		Figure 1. = Conversion of=20 Ink4a/Arf=96/=96 astrocytes to neural stem-like = cells by Id4.=20 (A) Up-regulation of neural stem cell markers in the=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes.=20 Immunofluorescence images (400x)=20 of GFAP (astrocyte marker), Nestin, Cd133, and Sox2 (neural = stem and=20 progenitor cell markers) in the vector-transduced=20 Ink4a/Arf=96/=96 astrocytes and Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes grown in DMEM + 10% = FBS culture=20 conditions. (B) Neurosphere formation of = Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes grown in neural stem = cell culture=20 conditions (DMEM/F12 + EGF [20 ng/mL] + bFGF [20 ng/mL]).=20 (Left panel) Representative photos showing = vector-transduced=20 Ink4a/Arf=96/=96 astrocytes and Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes grown in the neural = stem cell=20 culture for 7 d. (Right panel) Number of neurospheres = (10=9620=20 =B5m and >20 =B5m) of vector-transduced = Ink4a/Arf=96/=96=20 astrocytes and Id4-transduced Ink4a/Arf=96/=96 = astrocytes=20 (4000 cells per 24-well plate or one cell per square = millimeter)=20 grown in neural stem cell culture for 10 d. (*) P = < 0.01.=20 (C) Representative immunofluorescence images = (100x) showing Nestin and Cd133 in = the=20 cryomicrodissected neurospheres (10 =B5m) derived from = Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes. (D) = Neurosphere formation=20 during three serial passages of Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes (4000 cells per = 24-well plate)=20 grown in neural stem cell culture for 10 d. (*) P = < 0.01. =

 = ;

Tumor-initiating=20 frequency and histological characteristics of gliomas derived from=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes

Next, we assessed the tumorigenic potential of limiting cell=20 numbers following intracerebral injection and subsequent = detailed=20 clinical and histological assessment of injected brains. = While=20 no brain tumors formed with injection of 50,000 = vector-transduced=20 Ink4a/Arf=96/=96 astrocytes, the same number of = Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes formed high-grade = gliomas=20 within 3=964 wk (n =3D 5 mice) (Supplemental Fig. S4A).=20 Furthermore, three of three mice injected of 1000 = Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes developed tumors = within 5=966 wk=20 (data not shown), consistent with robust "tumor-initiating" = potential=20 as defined previously (Singh et al. 20043DGo). Upon histological examination, the tumors = derived=20 from Id4-transduced Ink4a/Arf=96/=96 cells = possessed hallmark=20 features of GBM including high cellularity, focal necrosis, = and=20 diffuse invasion (Fig. = 2A).=20 Thus, Id4 cooperates with Ink4a/Arf deficiency to enable = primary=20 astrocytes to attain a classical GBM phenotype.



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  		Figure 2. = Gliomagenesis=20 and in vivo differentiation of in the Id4 overexpressing=20 Ink4a/Arf=96/=96 astrocytes after intracerebral = implantation.=20 (A) No tumors (40x=20 magnification from Supplemental Fig. S4A, left photo) = observed in a nude mouse injected with = Ink4a/Arf=96/=96=20 astrocytes-Con-EGFP. Ink4a/Arf=96/=96=20 astrocytes-Id4-EGFP-derived gliomas (40x magnification from = Supplemental Fig.=20 S4A, right photo) showing a high cellularity, focal = necrosis, and=20 diffusive invasion. (B) Immunofluorescence images = (100x) showing Nestin+ = cells=20 (red) or Tuj1+ neuronal cells (green) or merge;=20 Nestin+ cells (red) or NG2+ = oligodendrocyte=20 cells (green) or merge; Nestin+ cells (red) or=20 CD31+ endothelial cells (green) or merge in brain = tissues=20 derived from orthotopically injected = Ink4a/Arf=96/=96=20 astrocytes-Id4-EGFP (brain tumor sections). Nuclei were = stained with=20 DAPI (blue). =

 = ;
Given=20 the role of Id4 in neuronal differentiation, we assessed the=20 differentiation profiles of cells or derivative tumors from=20 orthotopically injected Id4- and vector-transduced=20 Ink4a/Arf=96/=96 astrocytes which also express = EGFP. The=20 Id4-transduced Ink4a/Arf=96/=96 astrocyte-derived = tumors=20 showed many EGFP+ cells which stained positive for = Nestin=20 (NSC/progenitor), Tuj1 (neuronal), or NG2 (oligodendrocyte) = markers=20 and negative for the CD31 (endothelial cell) marker (Fig. = 2B for=20 tumors derived from injection of Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes; Supplemental Figs. = S4B for=20 EGFP, S5A for brain injected with vector-transduced = Ink4a/Arf=96/=96=20 astrocytes). None of the CD31+ cells expressed EGFP,=20 whereas all Nestin+, Tuj1+, or = NG2+=20 tumor cells were EGFP+ (data not shown), = indicating that=20 these cells are derived from the injected Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes. The same marker = profiles were=20 observed in tumor cells derived from subcutaneous injection = of=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes = (Cd133+,=20 Nestin+, or Tuj1+ cells) (Supplemental Fig. S5B).=20 Notably, most Tuj1+ cells did not express Cd133 or = Nestin,=20 and were closely associated with Cd133+ or=20 Nestin+ cells. In contrast to the Id4-transduced = cells,=20 the vector-transduced control cells failed to produce tumors = and=20 coimmunofluorescence using Tuj1/EGFP, NG2/EGFP, and GFAP/EGFP = antibodies did not show consistent colocalization patterns = for the=20 residual EGFP+ cells (data not shown). Thus, tumor = cells=20 derived from Id4-transduced Ink4a/Arf=96/=96 = astrocytes=20 express an array of lineage markers consistent with astrocyte = dedifferentation to an immature NSC/progenitor state with=20 multilineage potential and developmental plasticity.


Cyclin E contributes to elevated growth rate and = tumorigenicity of=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes

The exuberant growth rates associated with Id4 transduction=20 prompted analysis of cell cycle-positive and -negative = regulatory=20 factors, resulting in documentation of increased nuclear = cyclin=20 E and hyperphosphorylated Rb in Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes relative to controls = (Fig. = 3A=96C).=20 Next, gain- and loss-of-function studies revealed that cyclin = E=20 transduction increased proliferation in vector-transduced=20 Ink4a/Arf=96/=96 astrocytes (Fig. = 3D) to=20 rates comparable with those of Id4-transduced = Ink4a/Arf=96/=96=20 astrocytes (Supplemental Fig. S1D). Reciprocally, = shRNA-mediated=20 depletion of cyclin E caused markedly decreased proliferation = of=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes (Fig. = 3E).=20 These findings are in line with a prolonged G1/S profile in = the Id4=20 deficient embryonic dorsal telencephalon (Yun et al. 20043DGo). In tumor studies, cyclin E overexpression = in=20 Ink4a/Arf=96/=96 astrocytes resulted in tumor formation, = although these tumors were smaller in size than those = generated=20 from Id4-transduced Ink4a/Arf=96/=96 astrocytes (Fig.=20 3F). Conversely, shRNA-mediated depletion of cyclin E in=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes = resulted in=20 significant inhibition of tumor growth with an associated = decrease in=20 tumor cell proliferation as judged by PCNA staining (Fig. = 3F;=20 Supplemental Fig. S6A). Finally, a link between Id4 and = cyclin E was=20 further supported by colocalized expression in human GBM = specimens=20 (Supplemental Fig. S6B). Our cyclin E findings gain added=20 significance in light of previous studies establishing a = positive=20 correlation between increased cyclin E levels and poor = prognosis in=20 brain tumors (Tamiya et al. 20013DGo). Thus, oncogenic activity of Id4 appears to = be=20 executed in part through cyclin E-driven cell = proliferation.=20



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  		Figure 3. = Overexpression=20 of cyclin E in the Id4 overexpressing = Ink4a/Arf=96/=96=20 astrocytes is responsible for elevated growth rate and=20 tumorigenicity. (A) Expression of a variety of cell = cycle=20 regulators in the vector-transduced = Ink4a/Arf=96/=96=20 astrocytes and Id4-transduced Ink4a/Arf=96/=96 = astrocytes was=20 determined by Western blot analysis. (B) Expression = of cyclin=20 E protein in the vector-transduced = Ink4a/Arf=96/=96=20 astrocytes and Id4-transduced Ink4a/Arf=96/=96 = astrocytes was=20 determined by immunofluorescence. (C) Cyclin E mRNA = levels in=20 the vector-transduced Ink4a/Arf=96/=96 astrocytes = and=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes were = determined by=20 real-time RT=96PCR. (*) P < 0.05. (D) In = vitro growth=20 rate of vector-transduced Ink4a/Arf=96/=96 = astrocytes and=20 cyclin E-transduced Ink4a/Arf=96/=96 astrocytes.=20 (Inset) Expression levels of cyclin E in these cells = were=20 determined by Western blot analysis. (*) P < 0.01. = (E) In vitro growth inhibition of Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes that were subjected = to=20 down-regulation of cyclin E through cyclin E-specific shRNA = (cyclin=20 E RNAi). Expression levels of cyclin E in these cells were=20 determined by semiquantitative RT=96PCR (inset) and=20 immunofluorescence (right panel). (*) P < = 0.01.=20 (F) Decreased in vivo tumor growth via = down-regulation of=20 cyclin E in the Id4-transduced Ink4a/Arf=96/=96 = astrocytes.=20 Tumor weight (left panel) and representative photos=20 (right panel) showing nude mice injected = subcutaneously with=20 Ink4a/Arf=96/=96 astrocytes-Con, = Ink4a/Arf=96/=96=20 astrocytes-Id4, Ink4a/Arf=96/=96 = astrocytes-cyclin E, and=20 Ink4a/Arf=96/=96 astrocytes-Id4 + cyclin E-shRNA = (n =3D=20 6). (*) P < 0.01; (**) P < 0.01. =

 = ;

Id4=20 overexpression activates Notch signaling

Notch activation has been shown to be integral to stem cell=20 renewal (Chiba 20063DGo), the control of neuronal cell fate (Louvi = and=20 Artavanis-Tsakonas 20063DGo), and the malignant transformation of many = cancers=20 particularly lymphoid cells (Grabher et al. 20063DGo;=20 Hansson et al. 20043DGo). The dedifferentiation profile and NSC-like = phenotype=20 of Id4-transduced cells prompted us to examine components of = the=20 Notch signaling pathway (Artavanis-Tsakonas et al. 19993DGo). Relative to vector-transduced controls, Jagged1,=20 Presenilin-1, Hes1, Hey1, and Hey3 levels were significantly=20 increased in Id4-transduced Ink4a/Arf=96/=96 = astrocytes,=20 whereas Delta-like 1 and Hes6 expression levels were = comparable=20 (Fig.=20 4A). While Notch1 levels were unchanged, Id4 overexpressing = cells=20 showed markedly increased levels of activated Notch1 (NIC; = Notch=20 intracellular domain) (Fig. = 4B;=20 Supplemental Fig. S7A), as well as increased Notch/CSL = luciferase=20 reporter activity (Fig. = 4C).=20 The high levels of the Notch1 ligand, Jagged1, prompted = analysis of=20 Notch1 downstream signaling following shRNA-mediated = depletion of=20 Jagged1 in Id4-transduced Ink4a/Arf=96/=96 = astrocytes. Jagged1=20 knockdown resulted in decreased expression of NIC and its = downstream=20 targets, Hey1 and Hes1 (Fig.=20 4B). The relevance of Jagged1=96Notch signaling in GBM=20 pathogenesis is further supported by colocalization of Id4 = and Notch1=20 in human GBM specimens (Supplemental Fig. S7B).



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  		Figure 4. Notch = signaling=20 is activated by Id4, but is not associated with = tumorigenicity of=20 the Id4-transduced Ink4a/Arf=96/=96 astrocytes. = (A)=20 Expression levels of genes relative to Notch signaling were=20 determined by semiquantitative RT=96PCR. (*) P < = 0.01; (**)=20 P < 0.01. (B) Expression levels of Jagged1 = and=20 active Notch1 in the Ink4a/Arf=96/=96 = astrocytes-Con,=20 Ink4a/Arf=96/=96 astrocytes-Id4, and = Ink4a/Arf=96/=96=20 astrocytes-Id4 + Jagged1-shRNA were determined by Western = blot=20 analysis. 3D{alpha}=20-Tubulin was=20 used as a loading control. Expression levels of Hey1 and = Hes1=20 (Notch-downstream genes) in these cells were determined by=20 semiquantitative RT=96PCR. GAPDH was used as a loading = control.=20 (C) Relative Notch transcriptional activity in the=20 Ink4a/Arf=96/=96 astrocytes-Con, = Ink4a/Arf=96/=96=20 astrocytes-Id4, Ink4a/Arf=96/=96 astrocytes-Puro, = and=20 Ink4a/Arf=96/=96 astrocytes-NIC was determined by = Notch/CSL=20 luciferase-reporter assay. (*) P < 0.01. = (D) Cell=20 proliferation rate of Ink4a/Arf=96/=96 = astrocytes-Id4 and=20 Ink4a/Arf=96/=96 astrocytes-Id4 + Notch1-shRNA.=20 (Inset) Expression levels of Notch1 in these cells = were=20 determined by semiquantitative RT=96PCR. (*) P < = 0.01.=20 (E) Cell proliferation rates of = Ink4a/Arf=96/=96=20 astrocytes-Puro, and Ink4a/Arf=96/=96 = astrocytes-NIC.=20 (Inset) Expression levels of active NIC in these = cells were=20 determined by semiquantitative RT=96PCR. (F) = Representative=20 photos showing nude mice (n =3D 6) subcutaneously = injected with=20 Ink4a/Arf=96/=96 astrocytes-Con, = Ink4a/Arf=96/=96=20 astrocytes-Id4, Ink4a/Arf=96/=96 astrocytes-NIC, = and=20 Ink4a/Arf=96/=96 astrocytes-Id4 + Notch1-shRNA. =

 = ;
Next,=20 we assessed the pathogenetic contributions of activated = Notch1 to=20 Id4-induced gliomagenesis. shRNA-mediated Notch1 depletion in = Id4-transduced Ink4a/Arf=96/=96 astrocytes or NIC=20 overexpression in Ink4a/Arf=96/=96 astrocytes had = no=20 impact on proliferation rates (Fig. = 4D,E;=20 Supplemental Fig. S8). Furthermore, NIC overexpressing=20 Ink4a/Arf=96/=96 astrocytes were nontumorigenic = and,=20 reciprocally, Notch1 knockdown (80%) did not inhibit the = tumorigenic=20 potential of Id4-transduced Ink4a/Arf=96/=96 = astrocytes=20 (n =3D 6 mice) (Fig.=20 4F), suggesting that Notch1 does not play a direct role = in the=20 establishment of tumors from Id4-transduced = Ink4a/Arf=96/=96=20 astrocytes.


Id4-activated Notch signaling, but not cyclin E, is required = to=20 maintain the neural stem-like cell properties of the = Ink4a/Arf=96/=96=20 astrocytes

We next examined the impact of activated Notch signaling on = the=20 immature differentiation profiles and neurosphere-forming = capacity of=20 Id4-transduced Ink4a/Arf=96/=96 astrocytes. = Notch1, but not=20 cyclin E, knockdown resulted in a marked decrease in the = expression=20 of NSC markers, Nestin, Cd133, and Hes1, and correspondingly, = NIC=20 overexpression in Ink4a/Arf=96/=96 astrocytes = induced=20 expression of these immature markers (Fig.=20 5A). In the neurosphere assay, Notch1 knockdown resulted = in a=20 significant decrease in neurosphere number from the = Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes compared with a modest = decrease in=20 the cyclin E knockdown cultures (Fig. = 5B).=20 Furthermore, NIC overexpression, but not cyclin E, was = comparable=20 with Id4 in promoting neurosphere formation following their=20 transduction into Ink4a/Arf=96/=96 astrocytes (Fig. = 5C).=20 Pharmacological inhibition of Notch signaling (DAPT, a -secretase=20 inhibitor) or Jagged1 knockdown in Id4-transduced=20 Ink4a/Arf=96/=96 astrocytes resulted in decreased=20 neurosphere-forming capacity and NSC marker expression = (Supplemental=20 Fig. S9A=96C). Thus, Id4-induced activation of Jagged=96Notch = axis in=20 Ink4a/Arf=96/=96 astrocytes plays an essential = role in=20 promoting the neural stem cell-like phenotype.



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  		Figure 5. = Id4-activated=20 Notch signaling, but not cyclin E expression, is required to = maintain the neural stem-like cell properties of the=20 Ink4a/Arf=96/=96 astrocytes. (A) = Immunofluorescence=20 images (400x = magnification) of=20 Hes1 (Notch signaling marker), Nestin, and CD133 (neural = stem and=20 progenitor cell markers) in the Ink4a/Arf=96/=96=20 astrocytes-Con, Ink4a/Arf=96/=96 astrocytes-Id4,=20 Ink4a/Arf=96/=96 astrocytes-Id4 + Notch1-shRNA,=20 Ink4a/Arf=96/=96 astrocytes-Id4 + cyclin E-shRNA, = and=20 Ink4a/Arf=96/=96 astrocytes-NIC grown in DMEM + = 10% FBS=20 culture conditions. (B, left panel) Number of=20 neurospheres (>10 =B5m) generated from = Ink4a/Arf=96/=96=20 astrocytes-Con, Ink4a/Arf=96/=96 astrocytes-Id4,=20 Ink4a/Arf=96/=96 astrocytes-Id4 + cyclin E-shRNA, = and=20 Ink4a/Arf=96/=96 astrocytes-Id4 + Notch1-shRNA = (4000 cells=20 per 24-well plate) grown in neural stem cell cultures for 10 = d.=20 (Right panel) Representative photos (40x) showing the morphologies of = cells=20 grown in neural stem cell cultures. (*) P < 0.01.=20 (C, left panel) Number of neurospheres (>10 = =B5m)=20 generated from Ink4a/Arf=96/=96 astrocytes-Con,=20 Ink4a/Arf=96/=96 astrocytes-Id4, = Ink4a/Arf=96/=96=20 astrocytes-cyclin E, and Ink4a/Arf=96/=96 = astrocytes-NIC=20 (4000 cells per 24-well plate) grown in neural stem cell = cultures=20 for 10 d. (Right panel) Representative photos = (40x) showing the morphologies of = cells=20 grown in neural stem cell cultures. (*) P < 0.01. =

 = ;

Id4-dependent=20 Notch signaling and cyclin E expression in the human glioma cell = lines

Next, we sought to corroborate the murine findings in human = glioma=20 cells. shRNA-mediated depletion of Id4 in human LN229 glioma = cells=20 (which express the highest levels of Id4) (Supplemental Fig. = S1B)=20 resulted in down-regulation of cyclin E, Jagged1, NIC,=20 Notch-downstream target genes (Hes1, Hey1, and Hey2), and = Notch=20 transcriptional activity, as well as a marked decrease in = cell=20 proliferation (Supplemental Fig. S10A=96C). Conversely, Id4=20 overexpression in human A172 glioma cells (which express low=20 endogenous levels of Id4) induced up-regulation of Jagged1, = NIC, and=20 cyclin E; Notch transcriptional activity; cell proliferation; = and=20 neurosphere formation (Supplemental Fig. S10D=96F). We also = found that=20 expression levels of Id4, NIC, Jagged1, and cyclin E were = markedly=20 increased in the primary human glioma stem cell line, NCI0822 = (Lee et=20 al. 2008), as compared with NHA and HB1.F3 cells = (Supplemental=20 Fig. S10G).

Furthermore, we used a Tet-On-inducible gene expression system = to=20 assess whether Id4 directly leads to induction of cyclin E = and Notch=20 signaling. As shown in Supplemental Fig. S10H, Id4 was = markedly=20 increased in the Ink4a/Arf=96/=96 astrocytes = transduced with=20 rtTA and Rev-TRE-Id4 grown in the presence of doxycycline = (Dox) for 2=20 d compared with Dox-untreated counterpart cells. We also = found that=20 Dox-treated cells showed marked elevations in the levels of = cyclin E,=20 Jagged1, and NIC. These results strengthen the link between = Id4 and=20 the control of cyclin E and Notch signaling in the = astrocytes.=20

In conclusion, our findings that Id4 can drive the malignant=20 transformation of astrocytes via disregulation of cell cycle=20 and differentiation control, achieved through the = up-regulation=20 of cyclin E and activation of Jagged-Notch1 signaling. These=20 findings of Id4-induced developmental plasticity have = implications=20 for both the cellular origins of GBM as well as its prominent = renewal potential in experimental and clinical trials = setting.=20 Thus, these observations may inform the rational development = of=20 anti-Id4 agents that may impede the insuperable nature of GBM = recurrence.


3D"   =20 Materials and methods
Top
Abstract
3D"Results and Discussion
3D"Materials and methods
Acknowledgments
3D"References=20
 

Cell culture = and=20 conditions

Mouse astrocytes were isolated from the cerebral cortices of=20 5-d-old Ink4a/Arf=96/=96 knockout mice as = described=20 previously (Bachoo et al. 20023DGo). Ink4a/Arf=96/=96 astrocytes and = human GBM cell=20 lines were maintained in Dulbecco=92s modified Eagle=92s = medium (DMEM)=20 enriched with 10% fetal bovine serum (Hyclone), 1% = penicillin, and=20 streptomycin (GIBCO-BRL), and 2 mM L-glutamine (GIBCO-BRL). = For=20 neural sphere formation, cells (a single cell in each well of = a=20 96-well plate or 200 cells in a 24-well plate, which = corresponds to=20 one cell per square millimeter) were maintained in neural = stem cell=20 (NSC) culture conditions (DMEM/F12 medium enriched with 20 = ng/mL=20 EGF, 20 ng/mL bFGF, 1x = B27) and 1%=20 penicillin and streptomycin for 7=9610 d as described = previously, with=20 minor modification (Lee et al. 20063DGo).

Full materials and methods are available in the Supplemental=20 Material.


3D"   =20 Acknowledgments
Top
Abstract
3D"Results and Discussion
3D"Materials and methods
3D"Acknowledgments
3D"References =
 
We thank=20 Seung U. Kim for human neural stem cell (HB1.F3), Howard A. = Fine for=20 human glioma stem cell (NCI0822), and Eui-Ju Choi for the = Notch/CSL=20 luciferase=96reporter construct. This work was supported by a = grant=20 from the National R&D Program for Cancer Control, = Ministry of=20 Health and Welfare, Republic of Korea (0720030 to H.K.), and = NIH=20 grants (RO1CA99041 to L.C. and PO1CA95616 to R.A.D. and = L.C.). R.A.D.=20 is an American Cancer Society Research Professor and an = Ellison=20 Medical Foundation Scholar and is supported by the Robert A. = and=20 Renee E. Belfer Foundation Institute for Innovative Cancer=20 Science.


3D"   =20 Footnotes
 
5 Corresponding author. =

E-MAIL hg-kim{at}korea.ac.kr ; FAX 82-2-953-0737.

Supplemental material is available at = http://www.genesdev.org/. =

Article is online at http://www.g= enesdev.org/cgi/doi/10.1101/gad.1668708.=20


3D"   =20 References
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Abstract
3D"Results and Discussion
3D"Materials and methods
Acknowledgments
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Opera = requires=0A= the Sun Java Runtime Environment .=0A= =0A= by Andrew Gregory=0A= http://www.scss.com.au/family/andrew/webdesign/xmlhttprequest/=0A= =0A= This work is licensed under the Creative Commons Attribution License. To = view a=0A= copy of this license, visit = http://creativecommons.org/licenses/by-sa/2.5/ or=0A= send a letter to Creative Commons, 559 Nathan Abbott Way, Stanford, = California=0A= 94305, USA.=0A= =0A= Attribution: Leave my name and web address in this script intact.=0A= =0A= Not Supported in Opera=0A= ----------------------=0A= * user/password authentication=0A= * responseXML data member=0A= =0A= Not Fully Supported in Opera=0A= ----------------------------=0A= * async requests=0A= * abort()=0A= * getAllResponseHeaders(), getAllResponseHeader(header)=0A= =0A= */=0A= // IE support=0A= if (window.ActiveXObject && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= var msxmls =3D new Array(=0A= 'Msxml2.XMLHTTP.5.0',=0A= 'Msxml2.XMLHTTP.4.0',=0A= 'Msxml2.XMLHTTP.3.0',=0A= 'Msxml2.XMLHTTP',=0A= 'Microsoft.XMLHTTP');=0A= for (var i =3D 0; i < msxmls.length; i++) {=0A= try {=0A= return new ActiveXObject(msxmls[i]);=0A= } catch (e) {=0A= }=0A= }=0A= return null;=0A= };=0A= }=0A= // Gecko support=0A= /* ;-) */=0A= // Opera support=0A= if (window.opera && !window.XMLHttpRequest) {=0A= window.XMLHttpRequest =3D function() {=0A= this.readyState =3D 0; // = 0=3Duninitialized,1=3Dloading,2=3Dloaded,3=3Dinteractive,4=3Dcomplete=0A= this.status =3D 0; // HTTP status codes=0A= this.statusText =3D '';=0A= this._headers =3D [];=0A= this._aborted =3D false;=0A= this._async =3D true;=0A= this._defaultCharset =3D 'ISO-8859-1';=0A= this._getCharset =3D function() {=0A= var charset =3D _defaultCharset;=0A= var contentType =3D = this.getResponseHeader('Content-type').toUpperCase();=0A= val =3D contentType.indexOf('CHARSET=3D');=0A= if (val !=3D -1) {=0A= charset =3D contentType.substring(val);=0A= }=0A= val =3D charset.indexOf(';');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= val =3D charset.indexOf(',');=0A= if (val !=3D -1) {=0A= charset =3D charset.substring(0, val);=0A= }=0A= return charset;=0A= };=0A= this.abort =3D function() {=0A= this._aborted =3D true;=0A= };=0A= this.getAllResponseHeaders =3D function() {=0A= return this.getAllResponseHeader('*');=0A= };=0A= this.getAllResponseHeader =3D function(header) {=0A= var ret =3D '';=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= if (header =3D=3D '*' || this._headers[i].h =3D=3D header) {=0A= ret +=3D this._headers[i].h + ': ' + this._headers[i].v + '\n';=0A= }=0A= }=0A= return ret;=0A= };=0A= this.getResponseHeader =3D function(header) {=0A= var ret =3D getAllResponseHeader(header);=0A= var i =3D ret.indexOf('\n');=0A= if (i !=3D -1) {=0A= ret =3D ret.substring(0, i);=0A= }=0A= return ret;=0A= };=0A= this.setRequestHeader =3D function(header, value) {=0A= this._headers[this._headers.length] =3D {h:header, v:value};=0A= };=0A= this.open =3D function(method, url, async, user, password) {=0A= this.method =3D method;=0A= this.url =3D url;=0A= this._async =3D true;=0A= this._aborted =3D false;=0A= this._headers =3D [];=0A= if (arguments.length >=3D 3) {=0A= this._async =3D async;=0A= }=0A= if (arguments.length > 3) {=0A= opera.postError('XMLHttpRequest.open() - user/password not = supported');=0A= }=0A= this.readyState =3D 1;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= };=0A= this.send =3D function(data) {=0A= if (!navigator.javaEnabled()) {=0A= alert("XMLHttpRequest.send() - Java must be installed and = enabled.");=0A= return;=0A= }=0A= if (this._async) {=0A= setTimeout(this._sendasync, 0, this, data);=0A= // this is not really asynchronous and won't execute until the = current=0A= // execution context ends=0A= } else {=0A= this._sendsync(data);=0A= }=0A= }=0A= this._sendasync =3D function(req, data) {=0A= if (!req._aborted) {=0A= req._sendsync(data);=0A= }=0A= };=0A= this._sendsync =3D function(data) {=0A= this.readyState =3D 2;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= // open connection=0A= var url =3D new java.net.URL(new = java.net.URL(window.location.href), this.url);=0A= var conn =3D url.openConnection();=0A= for (var i =3D 0; i < this._headers.length; i++) {=0A= conn.setRequestProperty(this._headers[i].h, this._headers[i].v);=0A= }=0A= this._headers =3D [];=0A= if (this.method =3D=3D 'POST') {=0A= // POST data=0A= conn.setDoOutput(true);=0A= var wr =3D new = java.io.OutputStreamWriter(conn.getOutputStream(), this._getCharset());=0A= wr.write(data);=0A= wr.flush();=0A= wr.close();=0A= }=0A= // read response headers=0A= // NOTE: the getHeaderField() methods always return nulls for me :(=0A= var gotContentEncoding =3D false;=0A= var gotContentLength =3D false;=0A= var gotContentType =3D false;=0A= var gotDate =3D false;=0A= var gotExpiration =3D false;=0A= var gotLastModified =3D false;=0A= for (var i =3D 0; ; i++) {=0A= var hdrName =3D conn.getHeaderFieldKey(i);=0A= var hdrValue =3D conn.getHeaderField(i);=0A= if (hdrName =3D=3D null && hdrValue =3D=3D null) {=0A= break;=0A= }=0A= if (hdrName !=3D null) {=0A= this._headers[this._headers.length] =3D {h:hdrName, = v:hdrValue};=0A= switch (hdrName.toLowerCase()) {=0A= case 'content-encoding': gotContentEncoding =3D true; break;=0A= case 'content-length' : gotContentLength =3D true; break;=0A= case 'content-type' : gotContentType =3D true; break;=0A= case 'date' : gotDate =3D true; break;=0A= case 'expires' : gotExpiration =3D true; break;=0A= case 'last-modified' : gotLastModified =3D true; break;=0A= }=0A= }=0A= }=0A= // try to fill in any missing header information=0A= var val;=0A= val =3D conn.getContentEncoding();=0A= if (val !=3D null && !gotContentEncoding) = this._headers[this._headers.length] =3D {h:'Content-encoding', v:val};=0A= val =3D conn.getContentLength();=0A= if (val !=3D -1 && !gotContentLength) = this._headers[this._headers.length] =3D {h:'Content-length', v:val};=0A= val =3D conn.getContentType();=0A= if (val !=3D null && !gotContentType) = this._headers[this._headers.length] =3D {h:'Content-type', v:val};=0A= val =3D conn.getDate();=0A= if (val !=3D 0 && !gotDate) this._headers[this._headers.length] = =3D {h:'Date', v:(new Date(val)).toUTCString()};=0A= val =3D conn.getExpiration();=0A= if (val !=3D 0 && !gotExpiration) = this._headers[this._headers.length] =3D {h:'Expires', v:(new = Date(val)).toUTCString()};=0A= val =3D conn.getLastModified();=0A= if (val !=3D 0 && !gotLastModified) = this._headers[this._headers.length] =3D {h:'Last-modified', v:(new = Date(val)).toUTCString()};=0A= // read response data=0A= var reqdata =3D '';=0A= var stream =3D conn.getInputStream();=0A= if (stream) {=0A= var reader =3D new java.io.BufferedReader(new = java.io.InputStreamReader(stream, this._getCharset()));=0A= var line;=0A= while ((line =3D reader.readLine()) !=3D null) {=0A= if (this.readyState =3D=3D 2) {=0A= this.readyState =3D 3;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= }=0A= reqdata +=3D line + '\n';=0A= }=0A= reader.close();=0A= this.status =3D 200;=0A= this.statusText =3D 'OK';=0A= this.responseText =3D reqdata;=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onload) {=0A= this.onload();=0A= }=0A= } else {=0A= // error=0A= this.status =3D 404;=0A= this.statusText =3D 'Not Found';=0A= this.responseText =3D '';=0A= this.readyState =3D 4;=0A= if (this.onreadystatechange) {=0A= this.onreadystatechange();=0A= }=0A= if (this.onerror) {=0A= this.onerror();=0A= }=0A= }=0A= };=0A= };=0A= }=0A= // ActiveXObject emulation=0A= if (!window.ActiveXObject && window.XMLHttpRequest) {=0A= window.ActiveXObject =3D function(type) {=0A= switch (type.toLowerCase()) {=0A= case 'microsoft.xmlhttp':=0A= case 'msxml2.xmlhttp':=0A= case 'msxml2.xmlhttp.3.0':=0A= case 'msxml2.xmlhttp.4.0':=0A= case 'msxml2.xmlhttp.5.0':=0A= return new XMLHttpRequest();=0A= }=0A= return null;=0A= };=0A= }=0A= ------=_NextPart_000_0006_01C8FFBF.0FFB6830 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://genesdev.cshlp.org/javascript/ajax/utility.js /************************************************************************= *****=0A= * javascript/ajax/utility.js=0A= *=0A= * Utility functions for working with XMLHttpRequest data.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Copy XML nodes into an HTMLElement. This effectively=0A= * clones XML markup which uses XHTML naming conventions=0A= * into an HTML DOM.=0A= */=0A= function copy_xml_to_html(src, dst) {=0A= if (src.nodeType =3D=3D 1) { /* Node.ELEMENT_NODE */=0A= var e =3D document.createElement(src.nodeName);=0A= for (var i =3D 0; i < src.childNodes.length; i++) {=0A= copy_xml_to_html(src.childNodes[i], e);=0A= }=0A= for (var i =3D 0; i < src.attributes.length; i++) {=0A= var n =3D src.attributes[i].name;=0A= var v =3D unescape_xml_string(src.attributes[i].value); =0A= e.setAttribute(n, v);=0A= if (n =3D=3D "class") {=0A= e.className =3D v;=0A= }=0A= else if (n =3D=3D "style") {=0A= set_css_style(v, e, "");=0A= }=0A= }=0A= dst.appendChild(e);=0A= }=0A= else if (src.nodeType =3D=3D 3) { /* Node.TEXT_NODE */=0A= dst.appendChild(document.createTextNode(src.nodeValue));=0A= }=0A= }=0A= =0A= /* =0A= * It is unclear that this is the right thing to be calling=0A= * from copy_xml_to_html, but it appears that Safari decides=0A= * to convert & to the NCR #, and then encodes that=0A= * NCR to &%26%2338;. So, I'm going to treat the DOM Attr=0A= * value as a plain string, and run our XML string input=0A= * through the decoding routine below.=0A= */=0A= function unescape_xml_string(s) {=0A= return s.replace(/'/g, "'")=0A= .replace(/'/g, "'")=0A= .replace(/"/g, "\"")=0A= .replace(/"/g, "\"")=0A= .replace(/>/g, ">")=0A= .replace(/>/g, ">")=0A= .replace(/</g, "<")=0A= .replace(/</g, "<")=0A= .replace(/&/g, "&")=0A= .replace(/&/g, "&");=0A= }=0A= =0A= /*=0A= * Parse set of CSS rules and apply them to an element.=0A= * This is quite horrifying, but I'm unable to determine=0A= * how else to handle this with IE 6. FireFox and other=0A= * sane browsers let you simply set the style attribute=0A= * or use e.style.setProperty(rule, value, priority),=0A= * IE 6 appears to have neither of these capabilities..=0A= */=0A= function set_css_style(css, e, priority) {=0A= var rules =3D css.split(";");=0A= for (var i =3D 0; i < rules.length; i++) {=0A= var nvpair =3D rules[i].split(":");=0A= if (nvpair.length =3D=3D 2) {=0A= try {=0A= var name =3D nvpair[0]; /* style attribute */=0A= var value =3D nvpair[1]; /* attribute value */=0A= =0A= /*=0A= * For each possible style attribute, set the=0A= * appropriate style property in the element.=0A= */=0A= if (name =3D=3D "background") {=0A= e.style.background =3D value;=0A= }=0A= else if (name =3D=3D "background-attachment") {=0A= e.style.backgroundAttachment =3D value;=0A= }=0A= else if (name =3D=3D "background-color") {=0A= e.style.backgroundColor =3D value;=0A= }=0A= else if (name =3D=3D "background-image") {=0A= e.style.backgroundImage =3D value;=0A= }=0A= else if (name =3D=3D "background-position") {=0A= e.style.backgroundPosition =3D value;=0A= }=0A= else if (name =3D=3D "background-position-x") {=0A= e.style.backgroundPositionX =3D value;=0A= }=0A= else if (name =3D=3D "background-position-y") {=0A= e.style.backgroundPositionY =3D value;=0A= }=0A= else if (name =3D=3D "background-repeat") {=0A= e.style.backgroundRepeat =3D value;=0A= }=0A= else if (name =3D=3D "behavior") {=0A= e.style.behavior =3D value;=0A= }=0A= else if (name =3D=3D "border") {=0A= e.style.border =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom") {=0A= e.style.borderBottom =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-color") {=0A= e.style.borderBottomColor =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-style") {=0A= e.style.borderBottomStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-bottom-width") {=0A= e.style.borderBottomWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-collapse") {=0A= e.style.borderCollapse =3D value;=0A= }=0A= else if (name =3D=3D "border-color") {=0A= e.style.borderColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left") {=0A= e.style.borderLeft =3D value;=0A= }=0A= else if (name =3D=3D "border-left-color") {=0A= e.style.borderLeftColor =3D value;=0A= }=0A= else if (name =3D=3D "border-left-style") {=0A= e.style.borderLeftStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-left-width") {=0A= e.style.borderLeftWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-right") {=0A= e.style.borderRight =3D value;=0A= }=0A= else if (name =3D=3D "border-right-color") {=0A= e.style.borderRightColor =3D value;=0A= }=0A= else if (name =3D=3D "border-right-style") {=0A= e.style.borderRightStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-right-width") {=0A= e.style.borderRightWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-style") {=0A= e.style.borderStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top") {=0A= e.style.borderTop =3D value;=0A= }=0A= else if (name =3D=3D "border-top-color") {=0A= e.style.borderTopColor =3D value;=0A= }=0A= else if (name =3D=3D "border-top-style") {=0A= e.style.borderTopStyle =3D value;=0A= }=0A= else if (name =3D=3D "border-top-width") {=0A= e.style.borderTopWidth =3D value;=0A= }=0A= else if (name =3D=3D "border-width") {=0A= e.style.borderWidth =3D value;=0A= }=0A= else if (name =3D=3D "bottom") {=0A= e.style.bottom =3D value;=0A= }=0A= else if (name =3D=3D "clear") {=0A= e.style.clear =3D value;=0A= }=0A= else if (name =3D=3D "clip") {=0A= e.style.clip =3D value;=0A= }=0A= else if (name =3D=3D "color") {=0A= e.style.color =3D value;=0A= }=0A= else if (name =3D=3D "cssText") {=0A= e.style.Sets =3D value;=0A= }=0A= else if (name =3D=3D "cursor") {=0A= e.style.cursor =3D value;=0A= }=0A= else if (name =3D=3D "direction") {=0A= e.style.direction =3D value;=0A= }=0A= else if (name =3D=3D "display") {=0A= e.style.display =3D value;=0A= }=0A= else if (name =3D=3D "font") {=0A= e.style.font =3D value;=0A= }=0A= else if (name =3D=3D "font-family") {=0A= e.style.fontFamily =3D value;=0A= }=0A= else if (name =3D=3D "font-size") {=0A= e.style.fontSize =3D value;=0A= }=0A= else if (name =3D=3D "font-style") {=0A= e.style.fontStyle =3D value;=0A= }=0A= else if (name =3D=3D "font-variant") {=0A= e.style.fontVariant =3D value;=0A= }=0A= else if (name =3D=3D "font-weight") {=0A= e.style.fontWeight =3D value;=0A= }=0A= else if (name =3D=3D "height") {=0A= e.style.height =3D value;=0A= }=0A= else if (name =3D=3D "ime-mode") {=0A= e.style.imeMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-flow") {=0A= e.style.layoutFlow =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid") {=0A= e.style.layoutGrid =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-char") {=0A= e.style.layoutGridChar =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-line") {=0A= e.style.layoutGridLine =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-mode") {=0A= e.style.layoutGridMode =3D value;=0A= }=0A= else if (name =3D=3D "layout-grid-type") {=0A= e.style.layoutGridType =3D value;=0A= }=0A= else if (name =3D=3D "left") {=0A= e.style.left =3D value;=0A= }=0A= else if (name =3D=3D "letter-spacing") {=0A= e.style.letterSpacing =3D value;=0A= }=0A= else if (name =3D=3D "line-break") {=0A= e.style.lineBreak =3D value;=0A= }=0A= else if (name =3D=3D "line-height") {=0A= e.style.lineHeight =3D value;=0A= }=0A= else if (name =3D=3D "list-style") {=0A= e.style.listStyle =3D value;=0A= }=0A= else if (name =3D=3D "list-style-image") {=0A= e.style.listStyleImage =3D value;=0A= }=0A= else if (name =3D=3D "list-style-position") {=0A= e.style.listStylePosition =3D value;=0A= }=0A= else if (name =3D=3D "list-style-type") {=0A= e.style.listStyleType =3D value;=0A= }=0A= else if (name =3D=3D "margin") {=0A= e.style.margin =3D value;=0A= }=0A= else if (name =3D=3D "margin-bottom") {=0A= e.style.marginBottom =3D value;=0A= }=0A= else if (name =3D=3D "margin-left") {=0A= e.style.marginLeft =3D value;=0A= }=0A= else if (name =3D=3D "margin-right") {=0A= e.style.marginRight =3D value;=0A= }=0A= else if (name =3D=3D "margin-top") {=0A= e.style.marginTop =3D value;=0A= }=0A= else if (name =3D=3D "min-height") {=0A= e.style.minHeight =3D value;=0A= }=0A= else if (name =3D=3D "overflow") {=0A= e.style.overflow =3D value;=0A= }=0A= else if (name =3D=3D "overflow-x") {=0A= e.style.overflowX =3D value;=0A= }=0A= else if (name =3D=3D "overflow-y") {=0A= e.style.overflowY =3D value;=0A= }=0A= else if (name =3D=3D "padding") {=0A= e.style.padding =3D value;=0A= }=0A= else if (name =3D=3D "padding-bottom") {=0A= e.style.paddingBottom =3D value;=0A= }=0A= else if (name =3D=3D "padding-left") {=0A= e.style.paddingLeft =3D value;=0A= }=0A= else if (name =3D=3D "padding-right") {=0A= e.style.paddingRight =3D value;=0A= }=0A= else if (name =3D=3D "padding-top") {=0A= e.style.paddingTop =3D value;=0A= }=0A= else if (name =3D=3D "page-break-after") {=0A= e.style.pageBreakAfter =3D value;=0A= }=0A= else if (name =3D=3D "page-break-before") {=0A= e.style.pageBreakBefore =3D value;=0A= }=0A= else if (name =3D=3D "pixelBottom") {=0A= e.style.pixelBottom =3D value;=0A= }=0A= else if (name =3D=3D "pixelHeight") {=0A= e.style.pixelHeight =3D value;=0A= }=0A= else if (name =3D=3D "pixelLeft") {=0A= e.style.pixelLeft =3D value;=0A= }=0A= else if (name =3D=3D "pixelRight") {=0A= e.style.pixelRight =3D value;=0A= }=0A= else if (name =3D=3D "pixelTop") {=0A= e.style.pixelTop =3D value;=0A= }=0A= else if (name =3D=3D "pixelWidth") {=0A= e.style.pixelWidth =3D value;=0A= }=0A= else if (name =3D=3D "posBottom") {=0A= e.style.posBottom =3D value;=0A= }=0A= else if (name =3D=3D "posHeight") {=0A= e.style.posHeight =3D value;=0A= }=0A= else if (name =3D=3D "position") {=0A= e.style.position =3D value;=0A= }=0A= else if (name =3D=3D "posLeft") {=0A= e.style.posLeft =3D value;=0A= }=0A= else if (name =3D=3D "posRight") {=0A= e.style.posRight =3D value;=0A= }=0A= else if (name =3D=3D "posTop") {=0A= e.style.posTop =3D value;=0A= }=0A= else if (name =3D=3D "posWidth") {=0A= e.style.posWidth =3D value;=0A= }=0A= else if (name =3D=3D "right") {=0A= e.style.right =3D value;=0A= }=0A= else if (name =3D=3D "ruby-align") {=0A= e.style.rubyAlign =3D value;=0A= }=0A= else if (name =3D=3D "ruby-overhang") {=0A= e.style.rubyOverhang =3D value;=0A= }=0A= else if (name =3D=3D "ruby-position") {=0A= e.style.rubyPosition =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-3dlight-color") {=0A= e.style.scrollbar3dLightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-arrow-color") {=0A= e.style.scrollbarArrowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-base-color") {=0A= e.style.scrollbarBaseColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-darkshadow-color") {=0A= e.style.scrollbarDarkShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-face-color") {=0A= e.style.scrollbarFaceColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-highlight-color") {=0A= e.style.scrollbarHighlightColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-shadow-color") {=0A= e.style.scrollbarShadowColor =3D value;=0A= }=0A= else if (name =3D=3D "scrollbar-track-color") {=0A= e.style.scrollbarTrackColor =3D value;=0A= }=0A= else if (name =3D=3D "float") {=0A= e.style.styleFloat =3D value;=0A= }=0A= else if (name =3D=3D "table-layout") {=0A= e.style.tableLayout =3D value;=0A= }=0A= else if (name =3D=3D "text-align") {=0A= e.style.textAlign =3D value;=0A= }=0A= else if (name =3D=3D "text-align-last") {=0A= e.style.textAlignLast =3D value;=0A= }=0A= else if (name =3D=3D "text-autospace") {=0A= e.style.textAutospace =3D value;=0A= }=0A= else if (name =3D=3D "text-decoration") {=0A= e.style.textDecoration =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationBlink") {=0A= e.style.textDecorationBlink =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationLineThrough") {=0A= e.style.textDecorationLineThrough =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationNone") {=0A= e.style.textDecorationNone =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationOverline") {=0A= e.style.textDecorationOverline =3D value;=0A= }=0A= else if (name =3D=3D "textDecorationUnderline") {=0A= e.style.textDecorationUnderline =3D value;=0A= }=0A= else if (name =3D=3D "text-indent") {=0A= e.style.textIndent =3D value;=0A= }=0A= else if (name =3D=3D "text-justify") {=0A= e.style.textJustify =3D value;=0A= }=0A= else if (name =3D=3D "text-kashida-space") {=0A= e.style.textKashidaSpace =3D value;=0A= }=0A= else if (name =3D=3D "text-overflow") {=0A= e.style.textOverflow =3D value;=0A= }=0A= else if (name =3D=3D "text-transform") {=0A= e.style.textTransform =3D value;=0A= }=0A= else if (name =3D=3D "text-underline-position") {=0A= e.style.textUnderlinePosition =3D value;=0A= }=0A= else if (name =3D=3D "top") {=0A= e.style.top =3D value;=0A= }=0A= else if (name =3D=3D "unicode-bidi") {=0A= e.style.unicodeBidi =3D value;=0A= }=0A= else if (name =3D=3D "vertical-align") {=0A= e.style.verticalAlign =3D value;=0A= }=0A= else if (name =3D=3D "visibility") {=0A= e.style.visibility =3D value;=0A= }=0A= else if (name =3D=3D "white-space") {=0A= e.style.whiteSpace =3D value;=0A= }=0A= else if (name =3D=3D "width") {=0A= e.style.width =3D value;=0A= }=0A= else if (name =3D=3D "word-break") {=0A= e.style.wordBreak =3D value;=0A= }=0A= else if (name =3D=3D "word-spacing") {=0A= e.style.wordSpacing =3D value;=0A= }=0A= else if (name =3D=3D "word-wrap") {=0A= e.style.wordWrap =3D value;=0A= }=0A= else if (name =3D=3D "writing-mode") {=0A= e.style.writingMode =3D value;=0A= }=0A= else if (name =3D=3D "z-index") {=0A= e.style.zIndex =3D value;=0A= }=0A= else if (name =3D=3D "zoom") {=0A= e.style.zoom =3D value;=0A= }=0A= }=0A= catch (e) {=0A= /* ignore error on attempt to set e.style.[property] */=0A= }=0A= }=0A= }=0A= }=0A= ------=_NextPart_000_0006_01C8FFBF.0FFB6830 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://genesdev.cshlp.org/javascript/entrez/callback.js /************************************************************************= *****=0A= * javascript/entrez/callback.js=0A= *=0A= * Entrez Linking callback to populate content box.=0A= *=0A= * Copyright 2006 Board of Trustees of the Leland Stanford Junior = University.=0A= = *************************************************************************= ***/=0A= =0A= /*=0A= * Execute callback to fill content box with Entrez Linking information.=0A= */=0A= function entrez_callback(pmid, callback_url) {=0A= /*=0A= * MSIE 5.5 and below have issues with the JavaScript=0A= * used for Entrez Linking. 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