10.1007/s00401-009-0494-3</TITLE= > <META content=3D"text/html; charset=3DISO-8859-1" = http-equiv=3DContent-Type><LINK=20 rel=3DstyleSheet type=3Dtext/css=20 href=3D"http://www.springerlink.com/config/css/A++_common.css"><LINK=20 rel=3DstyleSheet type=3Dtext/css=20 href=3D"http://www.springerlink.com/config/css/content_A++_SpringerLink.c= ss"> <META name=3DGENERATOR content=3D"MSHTML 8.00.6001.18372"></HEAD> <BODY class=3Dfor-print> <TABLE border=3D1 cellPadding=3D2> <TBODY> <TR class=3Dheader> <TD>Acta Neuropathologica</TD></TR> <TR> <TD>=A9 Springer-Verlag 2009</TD></TR> <TR> <TD>10.1007/s00401-009-0494-3</TD></TR></TBODY></TABLE> <H2 class=3Drubric>Original Paper</H2> <DIV class=3DHeading1><A name=3Dtitle></A>Stem-cell-like glioma cells = are resistant=20 to TRAIL/Apo2L and exhibit down-regulation of caspase-8 by promoter = methylation=20 </DIV> David Capper1,=20 Timo Gaiser1, Christian Hartmann1, 3,=20 Antje Habel1, Wolf Mueller1,=20 Christel Herold-Mende2, = Andreas von Deimling1,=20 3 and Markus David Siegelin1 <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#ContactOfAuthor8"><IMG=20 border=3D0 alt=3D"Contact Information"=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/contact.gif">= </A> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff1></A>(1) </TD> <TD>Department of Neuropathology, = University=20 Hospital Heidelberg, Im Neuenheimer Feld 220, = 69120 Heidelberg,=20 Germany</TD></TR></TBODY></TABLE> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff2></A>(2) </TD> <TD>Department of Neurosurgery, University = Hospital Heidelberg, Im Neuenheimer Feld 400, = 69120 Heidelberg,=20 Germany</TD></TR></TBODY></TABLE> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff3></A>(3) </TD> <TD>Clinical Cooperation Unit = Neuropathology,=20 German Cancer Center (DKFZ) G380, Im Neuenheimer Feld 220,=20 69120 Heidelberg, Germany</TD></TR></TBODY></TABLE> <A name=3DContactOfAuthor8></A> <TABLE class=3DContact> <TBODY> <TR> <TD vAlign=3Dtop><IMG border=3D0 alt=3D"Contact Information"=20 = src=3D"http://www.springerlink.com/content/p536n18xr82w5576/contact.gif">= </TD> = <TD>Markus David Sieg= elin Email:=20 <A=20 = href=3D"mailto:markus.siegelin@med.uni-heidelberg.de">markus.siegelin@med= .uni-heidelberg.de</A></TD></TR></TBODY></TABLE> Received:=20 18 November 2008 Revised:=20 1 February 2009 Accepted:=20 1 February 2009 Published online:=20 12 February 2009 <DIV class=3DAbstract><A name=3DAbs1></A><SPAN=20 class=3DAbstractHeading>Abstract Tumour necrosis = factor=20 (TNF)-related apoptosis-inducing ligand (TRAIL/Apo2L) is a promising = cancer=20 drug. However, many tumours are resistant to TRAIL-based therapies. = Glioma cells=20 with stem cell features (SCG), such as CD133 expression and neurosphere=20 formation, have been recently identified to be more resistant to = cytotoxic drugs=20 than glioma cells lacking stem-cell-like features (NSCGs). Here we = report that=20 SCGs are completely resistant to 100=962,000 ng/ml TRAIL, whereas = NSCGs=20 revealed a moderate sensitivity to TRAIL. We found that SCGs exhibited = only low=20 levels of caspase-8 mRNA and protein, known to be indispensable for=20 TRAIL-induced apoptosis. In addition, we detected hypermethylation of=20 CASP8 promoter in SCGs, whereas NSCGs exhibited a non-methylated=20 CASP8 promoter. Reexpression of caspase-8 by = 5-Aza-2′-deoxycytidine was=20 not sufficient to restore TRAIL sensitivity in SCGs cells, suggesting = that=20 additional factors cause TRAIL resistance in SCGs. Our data suggest that = therapy=20 with TRAIL, either as monotherapy or in combination with demethylating = agents,=20 is not effective in treating glioblastoma because SCGs are not targeted = by such=20 treatment. </DIV> <SPAN=20 class=3DKeywordHeading>Keywords Stem-cell-like glioma=20 cells - Caspase-8 - TRAIL - Apoptosis = - CD133=20 <DIV class=3DKeyword><SPAN=20 class=3DKeywordHeading>Abbreviations <SPAN=20 class=3DTerm>IAP Inhibitor of apoptosis protein - <SPAN=20 class=3DTerm>TRAIL Tumour necrosis factor (TNF)-related=20 apoptosis-inducing ligand - SCG Stem-cell-like=20 glioma cells - NSCG Non-stem-cell-like = glioma cells=20 </DIV> <DIV><A name=3DSec1></A> <HR> <DIV class=3Dheading2>Introduction</DIV> Malignant gliomas are the most common malignant adult primary brain = tumours.=20 Median survival of glioblastoma patients is ~12=9615 months. = Current=20 treatment of malignant glioma usually comprises surgical resection or = diagnostic=20 biopsy followed by adjuvant radiation and chemotherapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR34">34</A></CITE>].=20 Recent studies reported on small sub-populations of tumour cells, = designated=20 as cancer stem cells (CSCs), differing from the majority of malignant = cells=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR32">32</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>].=20 These CSCs are thought to be more resistant to apoptosis, to survive = therapy,=20 and to eventually give rise to recurrent tumours, which usually are = highly=20 resistant to first-line therapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 The reasons for resistance of CSCs to therapy are not well known at = present,=20 although recent research indicates possible molecular determinants = underlying=20 this resistance, e.g., higher expression of BCRP1 and MGMT, as well as = the=20 anti-apoptosis protein and inhibitors of apoptosis protein families = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 Several proteins have been suggested as markers of CSCs, of which CD133 = has=20 received considerable attention because of its relatively high = expression in=20 CSCs of a variety of tumour types, including medulloblastomas [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>],=20 glioblastomas [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>],=20 colorectal [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR28">28</A></CITE>]=20 and prostate cancer [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR5">5</A></CITE>].=20 It has been shown that CD133+ cells, following their separation from = cultured=20 cancer cells or primary tumours, can give rise to glioblastomas in=20 immunocompromised (NOD/SCID) mice, which are phenotypically similar to = primary=20 tumours [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>].=20 It was reported that as many as 107 CD133-cultured = glioblastoma cells=20 were required to form tumours in NOD/SCID mice, whilst as few as = 103=20 CD133+ cells formed tumours in the same model [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR32">32</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>].=20 Tumour necrosis factor (TNF)-α-related apoptosis-inducing ligand = (TRAIL)=20 belongs to the TNF cytokine family and is capable of inducing apoptosis = in a=20 variety of cancer cells, whilst producing negligible effects on normal = cells=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR11">11</A></CITE>].=20 TRAIL binds to the death receptors DR4/DR5, which subsequently interact = with the=20 adaptor protein FADD and procaspase-8, forming the death-inducing = signalling=20 complex (DISC). Procaspase-8 activation in the DISC leads to cleavage of = procaspase-3 and engagement of the cellular machinery associated with = the type I=20 extrinsic apoptotic pathway [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR4">4</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR27">27</A></CITE>].=20 Activation of the intrinsic, mitochondrial-associated type II apoptotic = pathway=20 is another hallmark of TRAIL-induced cell death because TRAIL, through=20 caspase-8, activates Bid and synergizes with agents that induce = apoptosis=20 exclusively through a type II mechanism [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR6">6</A></CITE>].=20 In type I cells, stimulation of the extrinsic pathway is sufficient for=20 commitment of apoptotic cell death. In type II cells this commitment = requires=20 further signal amplification through the intrinsic pathway [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR23">23</A></CITE>].=20 Caspase-8 resides in the cytosol as a catalytically inactive zymogen and = is=20 present in two isoforms (p55, p53) [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR31">31</A></CITE>].=20 Caspase-8 zymogens are recruited by FADD to the DISC where they are = cleaved=20 through autoproteolysis, releasing the catalytically active p18 and p10 = subunits=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR31">31</A></CITE>].=20 The caspase-8 cleavage can be inhibited by cellular FADD-like=20 interleukin-1β-converting enzyme-inhibitory protein (c-FLIP) = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR17">17</A></CITE>].=20 There are two isoforms of c-FLIP: a short form (c-FLIPS) and = a long=20 form (c-FLIPL). Both c-FLIPS and = c-FLIPL are=20 recruited to the DISC where c-FLIPL is cleaved into a p43=20 intermediate form, which, together with c-FLIPS, remains in = the DISC=20 where they inhibit caspase-8 cleavage [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR39">39</A></CITE>].=20 Epigenetic alterations in DNA have been shown to be important in the = genesis=20 and progression of tumours, especially the methylation-mediated = silencing of=20 tumour suppressor genes [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR18">18</A></CITE>].=20 In cancer cells, aberrant methylation of CpG islands located in the = promoter=20 regions of genes implicated in apoptosis, migration, or DNA repair is = frequently=20 associated with transcriptional silencing and gene repression [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR18">18</A></CITE>].=20 The former alterations of the =91epigenome=92 also contribute to = defining the=20 biological behaviour of the tumour and might modulate the response of = tumour=20 cells to anticancer therapies [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR18">18</A></CITE>].=20 In the present study we found that CD133+ stem-cell-like glioblastoma = cells=20 (SCG) were completely resistant towards treatment with TRAIL and = exhibited only=20 low levels of caspase-8 mRNA and protein compared to CD133− = non-stem-cell-like=20 glioblastoma cells (NSCG). The up-regulation of caspase-8 by=20 5-Aza-2′-deoxycytidine treatment was not sufficient to sensitise = SCG to TRAIL,=20 indicating the existence of further factors mediating TRAIL resistance = in SCG.=20 </DIV> <DIV><A name=3DSec2></A> <HR> <DIV class=3Dheading2>Materials and methods</DIV> <DIV><A name=3DSec3></A> <DIV class=3DHeading3>Cell culture and reagents</DIV> Two paired cultures, NCH421K_SCG/NCH421K_NSCG and = NCH441_SCG/NCH441_NSCG,=20 were derived from surgical glioblastoma samples. The generation and=20 characteristics of these cell lines are described in detail elsewhere. = In brief,=20 glioma cell cultures derived from surgically removed tumour tissue were = FACS=20 sorted (see below) for CD133 expression and separated in CD133+ (SCG) = and CD133−=20 (NSCG) tumour cell fractions and were either established in foetal calf=20 serum-containing medium (NSCG) or a medium optimised for the growth of = normal=20 neural stem cells containing bFGF, EGF and the serum-free supplement = BIT9500=20 (SCG). SCG cells displayed stem cell typical growth as neurospheres, = increased=20 expression of CD133 and nestin, and were highly tumorigenic after = implantation=20 in NOD/SCID mice; 103 SCGs were sufficient to form tumours in = nude=20 mice, whereas 106 NSCG cells were necessary to achieve tumour = growth=20 in vivo [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR35">35</A></CITE>].=20 Common origin of the paired cell lines was confirmed by microsatellite = analysis=20 targeting polymorphisms on chromosomes 1 and 19 as previously described=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR13">13</A></CITE>].=20 NCH421K_NSCG and NCH441_NSCG were cultured in DMEM Glutamax-I = 4,500 g/l=20 glucose (Invitrogen, Karlsruhe, Germany) with 10% FBS and 1%=20 penicillin/streptomycin (Invitrogen, Karlsruhe) and were incubated at = 37=B0C in a=20 humidified atmosphere containing 10% carbon dioxide. NCH421_SCG and = NCH441_SCG=20 were incubated with serum-free neural stem cell medium containing = DMEM/F12=20 (Gibco), recombinant human epidermal growth factor (rhEGF, = 20 ng/ml; Sigma,=20 USA), basic fibroblast growth factor (bFGF, 20 ng/ml; Upstate, = USA), BIT:=20 bovine serum albumin, transferrin, insulin and 100 U/ml penicillin = G=20 (Provitro, Germany) and 100 μg/ml streptomycin (Gibco, USA). = Recombinant=20 human TRAIL/Apo2L was purchased from Peprotech (Rocky Hill, NY). = Cycloheximide=20 was obtained from Axxora (Loerrach, Germany), and cells were treated = with the=20 indicated amounts of cycloheximide as individually indicated. To exclude = the=20 effects of culture conditions on TRAIL resistance of SCG cells, SCG cell = lines=20 were additionally cultured for 48 h in DMEM Glutamax-I = 4,500 g/l=20 glucose with 10% FBS and 1% penicillin/streptomycin prior to TRAIL = treatment in=20 selected experiments without observable differences to culture = conditions in=20 neural stem cell medium. </DIV> <DIV><A name=3DSec4></A> <DIV class=3DHeading3>Caspase-3 activity assay</DIV> Cells were collected and analysed by Caspase-3 Colorimetric Assay Kit = (R&D Systems, Minneapolis, MN). Cells were lysed to collect their=20 intracellular contents. The cell lysate was tested for protease activity = by the=20 addition of a caspase-specific peptide that was conjugated to the colour = reporter molecule p-nitroanaline (pNA). The cleavage of the = peptide by=20 the caspase releases the chromophore pNA, which was quantitated=20 spectrophotometrically at a wavelength of 405 nm. The level of = caspase=20 enzymatic activity in the cell lysate was directly proportional to the = colour=20 reaction. </DIV> <DIV><A name=3DSec5></A> <DIV class=3DHeading3>Cell viability</DIV> Cells were seeded into 96-well plates at a density of=20 2 =D7 104 cells/well in 100-μl tissue = culture medium in=20 triplicate. After 24-h incubation allowing cells to adhere, cells were = treated=20 for 24 h either with cycloheximide or TRAIL or with combinations of = both,=20 as described in individual experiments. Cell viability was determined by = the=20 colorimetric = dimethylthiazolyl-carboxymethoxyphenyl-sulphophenyl-tetrazolium=20 (MTS) assay. In this assay, the quantity of formazan product formed is = directly=20 proportional to the number of viable cells in the cultures. After = 4 h in=20 culture the cell viability was determined by measuring the absorbance at = 490 nm using a 550 Bio-Rad plate-reader (Bio-Rad, Hertfordshire, = UK). The=20 relative percentage of survival was calculated by dividing the = absorbance of=20 treated cells by that of the control in each experiment. IC50 = values=20 (concentrations that led to a reduction of 50% cellular viability) were=20 calculated. </DIV> <DIV><A name=3DSec6></A> <DIV class=3DHeading3>Annexin V/PI staining</DIV> Apoptotic cells were assessed by annexin V staining using Annexin = V-FITC=20 apoptosis detection kit (BD Bioscience) following the manufacturer=92s=20 instructions and analysed by flow cytometry using Cellquest (Becton = Dickinson).=20 In the annexin V assay, the cells staining only for Annexin V-FITC = (early=20 apoptosis) are located in the lower right quadrant, whereas cells = staining for=20 both annexin V and propidiumiodide (PI) (late apoptosis) are located in = the=20 upper left quadrant. The percent positive cells for PI staining only in = the top=20 left quadrant represent the necrotic population. </DIV> <DIV><A name=3DSec7></A> <DIV class=3DHeading3>Flow cytometric analysis of protein expression and = cell=20 sorting</DIV> The level of expression of CD133 protein was assessed by flow = cytometry. In=20 brief, cells were harvested. They were then incubated with anti-CD133/2 = IgG=20 (clone 293C3; Miltenyi Biotec, Bergisch-Gladbach, Germany) followed by=20 incubation with secondary FITC- or PE-conjugated IgG (R&D Systems,=20 Wiesbaden, Germany). The cells were then sorted by flow cytometry = (FACSCalibur,=20 Becton Dickinson) for populations with high and low fluorescence. = </DIV> <DIV><A name=3DSec8></A> <DIV class=3DHeading3>Real-time reverse transcription PCR</DIV> Total RNA was isolated using the RNA Nucleospin RNA-II kit = (Macherey-Nagel,=20 D=FCren, Germany) according to the manufacturer=92s instructions. The = total RNA=20 concentration was assessed using a Nano-Drop photometer (Peqlab = Biotechnologie,=20 Erlangen, Germany). For cDNA synthesis 1 μg of total RNA was = used for=20 reverse transcription (random primer) using RevertAidTM H Minus MuLV = Polymerase=20 (1 U/μl), according to the manufacturer=92s instructions (MBI = Fermentas, St.=20 Leon-Rot, Germany). The following primer sequences were used: caspase-8 = forward,=20 5′CAG CAG CCT TGA AGG AAG TC3′; caspase-8 reverse, = 5′CGA GAT TGT CAT TAC CCC=20 ACA3′; c-FLIP forward, 5′CTC ACC GTC CCT GTA CCT3′; = c-FLIP reverse, 5′CAG GAG=20 TGG GCG TTT TCT3′; GAPDH forward, 5′TGC CTC TTT AGT TGT CAT = GCA G3′; GAPDH=20 reverse, 5′CCC GTT CAG CTC AGG GAT GA3′. All real-time PCR = reactions were=20 performed in a 25-μl mixture containing 1/20 volume of cDNA = preparation=20 (1 μl), 1=D7 SYBR Green buffer (PE Applied Biosystems, Foster = City, CA),=20 4 mM MgCl2, 0.2 μM of each primers (BA67 and = BA68),=20 0.2 mM dNTPs mix and 0.025 Unit of AmpliTaq Gold=AE = thermostable=20 DNA polymerase (Applied Biosystems, Foster City, CA). The PCR was run on = an ABI=20 PRISM Detection Instrument 7400. After 40 cycles, data reduction was = performed=20 with Sequence Detection System Software (Applied Biosystems Inc., Foster = City,=20 CA). For data analysis, threshold cycles (C T) for = GAPDH=20 (reference) and c-FLIP or caspase-8 (sample) were determined in = triplicates. We=20 chose cDNA of NCG421K_NSCG and NCH441_NSCG as calibrator cells (100%) = and=20 determined the relative change (rI) in copy numbers according to the = formula=20 rI =3D 2 exp[(C T c-FLIP or caspase-8=20 NSCG − C TGAPDH = NSCG) − (C=20 T c-FLIP or caspase-8 SCG − C = TGAPDH=20 SCG)]. </DIV> <DIV><A name=3DSec9></A> <DIV class=3DHeading3>Methylation-specific polymerase chain reaction = assay</DIV> DNA methylation patterns in the CpG islands of CASP8 were = determined=20 by chemical modification of unmethylated, but not the methylated, = cytosines to=20 uracil and subsequent polymerase chain reaction using primers specific = for=20 either methylated or the modified unmethylated DNA [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR8">8</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR14">14</A></CITE>].=20 One microgram of DNA was bisulphite treated and purified with the = EpiTect=20 Bisulfite Kit (QIAGEN Inc., Valencia, CA) following the manufacturer=92s = instructions. Primer sequences for methylation analysis are as described = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR26">26</A></CITE>]=20 (unmethylated sense: 5′-GTT GGT TTT ATT TAG TTT GGT-3′; = unmethylated antisense:=20 5′-CCC TAT CAA TAA CAA ATA ATA TAC-3′; methylated sense: = 5′-GTT GGT TTT ATT TAG=20 TTC GGC-3′; methylated antisense: 5′-CCC TAT CGA TAA CAA ATA = ATA TAC-3′).=20 Placental DNA treated in vitro with Sss I methyltransferase (New England = BioLabs, Beverly, MA) was used as positive control for methylated = alleles, and=20 DNA from normal lymphocytes was used as negative control for methylated = alleles.=20 Controls without DNA were performed for each set of polymerase chain = reaction.=20 Ten microlitres of each polymerase chain reaction was directly loaded = onto=20 non-denaturing 2% agarose gels, stained with ethidium bromide and = visualised=20 under UV illumination. </DIV> <DIV><A name=3DSec10></A> <DIV class=3DHeading3>Western blot</DIV> Twenty micrograms of protein diluted in NuPAGE-sample buffer and = reducing=20 reagent (Invitrogen, Carlsbad, Germany) were denatured at 95=B0C for = 5 min=20 and electrophoretically separated on ready-to-use 4=9612% sodium dodecyl = sulphate=96polyacrylamide gel electrophoresis (SDS=96PAGE) (Invitrogen, = Carlsbad).=20 Proteins were blotted onto nitrocellulose membranes at=20 1.5 mA/cm2 for 1.5 h (Invitrogen, Carlsbad). After = blocking=20 in 0.5 M Tris=96Base, pH 7.4, 5% milk powder, 1.5 M NaCl and = 0.05%=20 Tween, the membranes were incubated with mouse anti-human caspase-8 = antibody=20 diluted 1:1,000 (clone 1C12; CST, Danvers, MA), rabbit anti-human PARP = (clone=20 46D11; CST, Danvers), rabbit anti-human c-FLIP (R&D Systems, = Minneapolis,=20 MN), rabbit anti-cleaved caspase-3 (CST, Danvers) and rabbit anti-human=20 caspase-7 (clone Asp198; CST Inc., Danvers) overnight at 4=B0C. Staining = with=20 secondary horseradish peroxidase conjugated anti-rabbit or anti-mouse = antibodies=20 at dilutions of 1:10,000 or 1:2,000, respectively (Amersham Biosciences, = Buckinghamshire, UK) was followed by immunodetection with Western = Blotting=20 Detection System ECL (Amersham Biosciences, Buckinghamshire). Protein = signals=20 were analysed semiquantitatively, using a computer-assisted image = analysis=20 system and the NIH gel analysis software (<A=20 href=3D"http://www.rsb.info.nih.gov/nihimage/download.html">http://www.rs= b.info.nih.gov/nihimage/download.html</A>).=20 </DIV> <DIV><A name=3DSec11></A> <DIV class=3DHeading3>Human Apoptosis Array Kit/Proteome = Profiler=99</DIV> For analysing the expression profiles of apoptosis-related proteins, = we used=20 the Human Apoptosis Array Kit (R&D Systems Ltd., Abingdon, UK). This = assay=20 consists of 30 nitrocellulose membrane spotted antibodies specific for=20 apoptosis-related proteins. Cell samples (1 =D7 107 = cells)=20 were harvested, and 300 μg of protein was mixed with = 15 μl of=20 biotinylated detection antibodies. After pre-treatment the cocktail was=20 incubated overnight at 4=B0C on a rocking platform. Following a wash = step to=20 remove unbound material, streptavidin=96horseradish and chemiluminescent = detection=20 reagents were added sequentially. Chemiluminescence at each spot is = proportional=20 to the amount of bound substrate. All experiments were repeated three = times. The=20 signals on X-ray film were quantified by scanning on a transmission-mode = scanner=20 and analysing the array image file using ImageJ analysis software (<A=20 href=3D"http://rsbweb.nih.gov/ij/">http://rsbweb.nih.gov/ij/</A>). In = detail, the=20 following procedure was carried out: The averaged background signal was subtracted and then the arrays = were=20 calibrated based on the signal strength of the positive controls. The = average=20 signal (pixel density) of the pair of duplicated spots representing each = apoptosis protein was determined. The corresponding signals on different = arrays=20 were compared and the relative change in apoptosis proteins was = determined.=20 </DIV></DIV> <DIV><A name=3DSec12></A> <HR> <DIV class=3Dheading2>Results</DIV> <DIV><A name=3DSec13></A> <DIV class=3DHeading3>SCGs express CD133</DIV> <DIV class=3DPara> <DIV>NCH421K_SCG, NCH421K_NSCG, NCH441_SCG and NCH441_NSCG were analysed = for the=20 expression of CD133 by flow cytometry. Two representative flow cytometry = charts=20 are shown (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig1">1</A>a,=20 b). In stem-cell-like glioma cell culture, 91% of NCH421K_SCG and 92% = NCH441_SCG=20 stained positive for CD133 (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig1">1</A>c).=20 In contrast, NCH421K_NSCG and NCH441_NSCG cultures contained only 1 and = 2% CD133=20 positive cells, respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig1">1</A>c).=20 <DIV class=3DFigure><A name=3DFig1></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig1_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig1_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 1 Rate of = CD133 in SCGs=20 and NSCGs. Flow cytometry of NCH421K_SCG (a) and NCH441_SCG = (b)=20 stained with CD133 antibody. M2 represents cells positive, whereas M1 = shows=20 cells negative for CD133. c Diagram of CD133 rate in NCH421K_SCG, = NCH441_SCG, NCH421K_NSCG and NCH441_NSCG determined by flow cytometry = </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec14></A> <DIV class=3DHeading3>SCGs are resistant towards TRAIL treatment</DIV> <DIV class=3DPara> <DIV>SCGs were resistant to TRAIL treatment (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>).=20 Treatment with 100, 250, 500, 1,000 and 2,000 ng/ml TRAIL for = 24 h=20 alone did not have a significant effect on cell death in NCH441_SCG=20 (IC50 > 2,000 ng/ml) and NCH421K_SCG=20 (IC50 > 2,000 ng/ml) (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>a).=20 Even prolonged treatment with TRAIL for up to 7 days did not induce = cell=20 death in SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>b).=20 In contrast, NSCGs were moderately sensitive to TRAIL. Treatment with = different=20 concentrations of TRAIL for 24 h had a significant effect on cell = death in=20 NCH441_NSCG and NCH421_NSCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>a).=20 The IC50s were calculated for NCH441_NSCG=20 (220 =B1 40 ng/ml) and NCH421K_NSCG=20 (177 =B1 52 ng/ml).=20 <DIV class=3DFigure><A name=3DFig2></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig2_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig2_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 2 MTT = viability assay=20 and Western blot analysis after treatment with TRAIL, cycloheximide or = the=20 combination of both after 24 h of treatment. a MTT assay in=20 NCH421K_SCG, NCH441_SCG, NCH421K_NSCG and NCH441_NSCG cells upon = treatment with=20 TRAIL for 24 h. b MTT assay in NCH421K_SCG and NCH441_SCG = upon=20 treatment with 1,000 ng/ml TRAIL for up to 7 days. c = MTT assay=20 in NCH421K_SCG, NCH441_SCG, NCH421K_NSCG and NCH441_NSCG cells upon = treatment=20 with TRAIL, cycloheximide or the combination of both for 24 h. = d=20 Western blot showing cleaved caspase-7 and PARP in NCH421K_SCG, = NCH441_SCG,=20 NCH421K_NSCG and NCH441_NSCG untreated or treated with 500 ng/ml = TRAIL for=20 8 h. e Caspase-3 activity in NCH421K_SCG, NCH441_SCG, = NCH421K_NSCG=20 and NCH441_NSCG upon treatment with TRAIL. Control not treated,=20 TR100/250/500/1000/2000 TRAIL 100/250/500/1,000/2,000 ng/ml, = 0d/1d/2d/5d/7d day 0/1/2/5/7 after start of treatment with TRAIL, = CHX20 cycloheximide 20 μg/ml. Experiments were = performed at least=20 three times. Asterisks outline values that are different from the = respective control (t test, *P < 0.05); = n.s.=20 not significant </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec15></A> <DIV class=3DHeading3>Inhibition of protein synthesis by cycloheximide = does not=20 sensitise SCGs to TRAIL-mediated cytotoxicity, but enhances = TRAIL-induced=20 cytotoxicity in NSCGs </DIV> After 24 h treatment with cycloheximide 20 μg/ml cell = death=20 increased significantly in NCH441_SCG (27 =B1 3%) and = NCH421K_SCG=20 (36 =B1 3%), respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Combining 20 μg/ml cycloheximide with 1,000 ng/ml of = TRAIL did not=20 increase cell death significantly in NCH441_SCG (30 =B1 3%) = and=20 NCH421K_SCG (44 =B1 3%) compared to single treatment with = cycloheximide=20 (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Thus, the protein synthesis inhibitor was not capable of sensitising = SCGs to=20 TRAIL-mediated cytotoxicity. Single treatment with 20 μg/ml cycloheximide increased cell = death in=20 NCH441_NSCG (18 =B1 3%) and NCH421K_NSCG = (17 =B1 3%),=20 respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Combining 20 μg/ml cycloheximide with 100 ng/ml of = TRAIL increased=20 cell death significantly in NCH441_NSCG (65 =B1 3%) and = NCH421K_NSCG=20 (74 =B1 3%) compared to single treatment with cycloheximide or = TRAIL-only treated cells (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Thus, in NSCGs the protein synthesis inhibitor further enhanced the = cytotoxic=20 effect of TRAIL. </DIV> <DIV><A name=3DSec16></A> <DIV class=3DHeading3>TRAIL mediates activation (cleavage) of = effector-caspases 3,=20 7 and poly-ADP-ribose-polymerase in NSCGs</DIV> We employed Western blotting to elucidate the proteolytic mechanism = in=20 TRAIL-induced apoptosis (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>d).=20 NCH441_SCG, NCH421K_SCG, NCH441_NSCG and NCH421K_NSCG cells were treated = with=20 TRAIL for 12 h, and the activation/cleavage of caspase-7 and=20 poly-ADP-ribose-polymerase (PARP) were examined. Exposure of cells to=20 500 ng/ml TRAIL alone yielded undetectable signals of cleaved = fragment of=20 PARP sized 89 kDa and active cleaved caspase-7 sized 20 kDa in = NCH441_SCG and NCH421K_SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>d).=20 Treatment with 500 ng/ml TRAIL led to a significant increase in = cleaved=20 fragment of PARP and active cleaved caspase-7/3 in NCH441_NSCG and = NCH421K_NSCG=20 (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>d).=20 Upon treatment with 500 or 1,000 ng/ml TRAIL SCG did not exhibit an = increase in caspase-3 activity, whereas NSCG showed a significant = increase in=20 caspase-3 activity compared to the controls (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>e).=20 </DIV> <DIV><A name=3DSec17></A> <DIV class=3DHeading3>Human Apoptosis Array Kit/Proteome = Profiler=99</DIV> <DIV class=3DPara> <DIV>To identify the resistance mechanism leading to TRAIL resistance in = SCGs we=20 analysed the expression of 30 apoptosis-related proteins that have = already been=20 shown to be associated with TRAIL resistance (Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>a;=20 Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig3">3</A>a,=20 b). Concentrations of the following proteins were significantly = different in SCG=20 and NSCG: Bad, cleaved caspase-3 Fas/TNFSF6, TRAIL R2/DR5, phospho-p53 = (S392),=20 XIAP, FADD, HSP60 and p21/CIP1/CDNK1A (Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>a,=20 b). DR5 and FADD were significantly increased in NCH441_NSCG compared to = NCH441_SCG, indicating a higher susceptibility of NCH441_NSCG to = TRAIL-mediated=20 apoptosis. However, only 2 of the 30 analysed proteins were = differentially=20 regulated in both cell line pairs. We detected a significant = up-regulation of=20 Fas/TNFSF6 and a down-regulation of phospho-p53 (S392) in NSCGs = (Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>b,=20 c).<A name=3DTab1></A> <DIV class=3DCapt>Table 1 Human = Apoptosis=20 Array Kit/Proteome Profiler=99 </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH colSpan=3D4 align=3Dleft> (a)</TH></TR> <TR class=3Dheader> <TH align=3Dleft> Coordinate</TH> <TH align=3Dleft> Target/control</TH> <TH align=3Dleft> Coordinate</TH> <TH align=3Dleft> Target/control</TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> A1, A2</TD> <TD align=3Dleft> Positive control</TD> <TD align=3Dleft> C13, C14</TD> <TD align=3Dleft> HO-2/HMOX2</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> A23, A24</TD> <TD align=3Dleft> Positive control</TD> <TD align=3Dleft> C15, C16</TD> <TD align=3Dleft> HSP27</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B1, B2</TD> <TD align=3Dleft> Bad</TD> <TD align=3Dleft> C17, C18</TD> <TD align=3Dleft> HSP60</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B3, B4</TD> <TD align=3Dleft> Bax</TD> <TD align=3Dleft> C19, C20</TD> <TD align=3Dleft> HSP70</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B5, B6</TD> <TD align=3Dleft> Bcl-2</TD> <TD align=3Dleft> C21, C22</TD> <TD align=3Dleft> HTRA2/Omi</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B7, B8</TD> <TD align=3Dleft> Bcl-x</TD> <TD align=3Dleft> C23, C24</TD> <TD align=3Dleft> Livin</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B9, B10</TD> <TD align=3Dleft> Procaspase-3</TD> <TD align=3Dleft> D1, D2</TD> <TD align=3Dleft> PON2</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B11, B12</TD> <TD align=3Dleft> Cleaved caspase-3</TD> <TD align=3Dleft> D3, D4</TD> <TD align=3Dleft> p21/CIP1/CDNK1A</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B13, B14</TD> <TD align=3Dleft> Catalase</TD> <TD align=3Dleft> D5, D6</TD> <TD align=3Dleft> p27/Kip1</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B15, B16</TD> <TD align=3Dleft> cIAP-1</TD> <TD align=3Dleft> D7, D8</TD> <TD align=3Dleft> Phospho-p53 (S15)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B17, B18</TD> <TD align=3Dleft> cIAP-2</TD> <TD align=3Dleft> D9, D10</TD> <TD align=3Dleft> Phospho-p53 (S46)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B19, B20</TD> <TD align=3Dleft> Claspin</TD> <TD align=3Dleft> D11, D12</TD> <TD align=3Dleft> Phospho-p53 (S392)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B21, B22</TD> <TD align=3Dleft> Clusterin</TD> <TD align=3Dleft> D13, D14</TD> <TD align=3Dleft> Phospho-Rad17 (S635)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B23, B24</TD> <TD align=3Dleft> Cytochrome c </TD> <TD align=3Dleft> D15, D16</TD> <TD align=3Dleft> SMAC/Diablo</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C1, C2</TD> <TD align=3Dleft> TRAIL R1/DR4</TD> <TD align=3Dleft> D17, D18</TD> <TD align=3Dleft> Survivin</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C3, C4</TD> <TD align=3Dleft> TRAIL R2/DR5</TD> <TD align=3Dleft> D19, D20</TD> <TD align=3Dleft> TNF RI/TNFRSF1A</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C5, C6</TD> <TD align=3Dleft> FADD</TD> <TD align=3Dleft> D21, D22</TD> <TD align=3Dleft> XIAP</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C7, C8</TD> <TD align=3Dleft> Fas/TNFSF6</TD> <TD align=3Dleft> D23, D24</TD> <TD align=3Dleft> PBS (negative control)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C9, C10</TD> <TD align=3Dleft> HIF-1</TD> <TD align=3Dleft> E1, E2</TD> <TD align=3Dleft> Positive control</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C11, C12</TD> <TD align=3Dleft> HO1/HMOX1/HSP32</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD></TR></TBODY></TABLE> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dchar char=3D"."> <COL align=3Dchar char=3D"=B1"> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH colSpan=3D4 align=3Dleft> (b)</TH></TR> <TR class=3Dheader> <TH align=3Dleft> </TH> <TH align=3Dleft char=3D"."> NCH421K_SCG</TH> <TH align=3Dleft char=3D"=B1"> NCH421K_NSCG</TH> <TH align=3Dleft> </TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> Bad </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.56 =B1 0.15 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bax</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.2 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-x</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.0 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Procaspase-3</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.26</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cleaved caspase-3 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 3.43 =B1 0.24 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Catalase</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.10</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Claspin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.15 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Clusterin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cytochrome c </TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R1/DR4</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.07 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R2/DR5</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.10</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> FADD</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Fas/TNFSF6 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 10.34 =B1 0.22 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HIF-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.16</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO1/HMOX1/HSP32</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO-2/HMOX2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.11 =B1 0.24</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP27</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.55 =B1 0.23</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP60</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.23 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP70</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.35 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HTRA2/Omi</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Livin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.13 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> PON2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.22 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p21/CIP1/CDNK1A</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.35 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p27/Kip1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.91 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S15)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.94 =B1 0.17</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S46)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S392) </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.55 =B1 0.21 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-Rad17 (S635)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.2 =B1 0.29</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> SMAC/Diablo</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.22</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Survivin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.27</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TNF RI/TNFRSF1A</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.28</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> XIAP </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.45 =B1 0.26 </TD> <TD align=3Dleft> </TD></TR></TBODY></TABLE> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dchar char=3D"."> <COL align=3Dchar char=3D"=B1"> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH colSpan=3D4 align=3Dleft> (c)</TH></TR> <TR class=3Dheader> <TH align=3Dleft> </TH> <TH align=3Dleft char=3D"."> NCH441_SCG</TH> <TH align=3Dleft char=3D"=B1"> NCH441_NSCG</TH> <TH align=3Dleft> </TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> Bad</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bax</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.28 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.15 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-x</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.0 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Procaspase-3</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.26</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cleaved caspase-3</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.24</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Catalase</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.10</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Claspin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.15 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Clusterin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cytochrome c </TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R1/DR4</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.23 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R2/DR5 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 1.88 =B1 0.15 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> FADD </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 1.82 =B1 0.18 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Fas/TNFSF6 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 16.12 =B1 0.22 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HIF-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.16</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO1/HMOX1/HSP32</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO-2/HMOX2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.11 =B1 0.24</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP27</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.55 =B1 0.23</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP60 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.56 =B1 0.13 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP70</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.35 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HTRA2/Omi</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.28</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Livin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.13 =B1 0.23</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> PON2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.22 =B1 0.25</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p21/CIP1/CDNK1A </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 8.56 =B1 0.54 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p27/Kip1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.91 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S15)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.94 =B1 0.17</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S46)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S392) </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.18 =B1 0.23 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-Rad17 (S635)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.2 =B1 0.29</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> SMAC/Diablo</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.22</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Survivin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.27</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TNF RI/TNFRSF1A</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.28</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> XIAP</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.13 =B1 0.26</TD> <TD align=3Dleft> </TD></TR></TBODY></TABLE> <DIV class=3DCapt> <DIV class=3DCaptCont> <DIV>a: Identification of the 30 apoptosis-related proteins on the = protein=20 array. b, c: Expression level of 30 apoptosis-related proteins in = NCH421K_SCG=20 (b), NCH441_SCG (c), NCH421_NSCG (b) and NCH441_NSCG (b). Protein levels = of SCG=20 were normalised to 1, and results are given as relative values with=20 corresponding standard deviations. Proteins and values in bold were=20 significantly different. Arrays and analysis were performed thrice=20 </DIV></DIV></DIV> <DIV class=3DFigure><A name=3DFig3></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig3_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig3_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 3 Analysis = of 30=20 apoptosis-related proteins in SCGs and NSCGs. a Stem-cell-like=20 (NCH421K_SCG) and non-stem-cell-like glioblastoma cells (NCH421K_NSCG) = were=20 harvested and incubated with nitrocellulose membranes containing 30=20 apoptosis-related proteins. b Stem-cell-like (NCH441_SCG) and=20 non-stem-cell-like glioblastoma cells (NCH441_NSCG) were harvested and = incubated=20 with nitrocellulose membranes containing 30 apoptosis-related proteins. = Proteins=20 with a significantly different expression are marked and outlined in the = figure.=20 For results of densitometry, refer to Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>=20 </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec18></A> <DIV class=3DHeading3>Caspase-8 expression is suppressed in SCGs</DIV> Since the human apoptosis array kit does not include c-FLIP and = caspase-8, we=20 analysed the expression of c-FLIP and caspase-8 by real-time reverse=20 transcription PCR and Western blot. <DIV class=3DPara> <DIV>Significantly different levels of c-FLIP mRNA or protein were not = observed=20 between SCGs and NSCGs (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig4">4</A>a=96c).=20 <DIV class=3DFigure><A name=3DFig4></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig4_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig4_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 4 Expression analysis=20 of c-FLIP and caspase-8 mRNA and protein level. a, b mRNA = levels=20 of c-FLIP and caspase-8 were analysed in NCH421K_SCG, NCH421K_NSCG, = NCH441_SCG=20 and NCH441_NSCG. The levels of c-FLIP and caspase-8 were normalised to 1 = in=20 non-stem-cell-like glioma cells. Experiments were repeated thrice. = c=20 Western blot analysis of c-FLIP and caspase-8 in NCH421K_SCG, = NCH421_NSCG,=20 NCH441_SCG and NCH441_NSCG. Actin served as protein loading control = </DIV> <HR> </DIV></DIV> Both SCG cell lines demonstrated significantly reduced levels of = caspase-8=20 compared to NSCGs. In NCH421K_SCG the relative mRNA level of caspase-8 = was=20 76 =B1 3% (P < 0.01) lower than in = NCH421K_NSCG, and=20 in NCH441_SCG it was 98 =B1 5% (P < 0.01) = lower than=20 in NCH441_NSCG, respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig4">4</A>a,=20 b). These results were confirmed on the protein level. The protein = levels of=20 caspase-8 were significantly reduced in both SCGs, NC441_SCG and = NCH421K_SCG,=20 compared to the NSCG cells (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig4">4</A>c).=20 </DIV> <DIV><A name=3DSec19></A> <DIV class=3DHeading3>Hypermethylation of the CASP8 promoter = region in SCGs=20 </DIV> <DIV class=3DPara> <DIV>Caspase-8 was suppressed at the level of transcription. Therefore,=20 silencing of caspase-8 in SCG might be due to methylation of the = CASP8=20 promoter region. We employed methylation-specific polymerase chain = reaction to=20 assess the methylation status of CASP8 as described in the = materials and=20 methods section. Both SCGs, NCH421K_SCG and NCH441_SCG, demonstrated a=20 methylated CASP8 promoter (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>a).=20 In contrast, NSCGs cells exhibited no methylation of the CASP8 = promoter=20 in both cell lines NCH421K_NSCG and NCH441_NSCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>a).=20 Importantly, treatment with 5-Aza-2′-deoxycytidine resulted in a = significant=20 up-regulation of caspase-8 on both mRNA and protein levels in NCH421_SCG = and=20 NCH441_SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>b,=20 c), suggesting that hypermethylation of CASP8 promoter is = responsible for=20 suppression of caspase-8 levels in SCG cells.=20 <DIV class=3DFigure><A name=3DFig5></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig5_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig5_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 5 Analysis = of CpG=20 island promoter methylation status in SCGs and NSCGs by the = methylation-specific=20 polymerase chain reaction assay. a The presence of a visible = polymerase=20 chain reaction product in those lanes marked U indicates the = presence of=20 an unmethylated CASP8 promoter; the presence of product in those = lanes=20 marked M indicates the presence of a methylated CASP8 = promoter.=20 Water (H2O) was used as negative polymerase chain reaction = control.=20 In vitro methylated placental DNA (IVD) was used as positive control for = methylated genes. DNA from normal lymphocytes (NLs) was used as negative = control=20 for methylated alleles. b mRNA levels of caspase-8 NCH421_SCG and = NCH441_SCG after treatment with different concentrations (μM) of=20 5-Aza-2′-deoxycytidine (5 Aza) for 120 h. c Protein = levels of=20 caspase-8 after treatment with different concentrations (μM) of=20 5-Aza-2′-deoxycytidine (5 Aza) for 120 h. LN229 is an = established cell line=20 expressing considerable levels of caspase-8. d Annexin V/PI flow=20 cytometric based assay in NCH421K_SCG, NCH441_SCG, upon treatment with = TRAIL,=20 5-Aza-2′-deoxycytidine or the combination of both for 72 h. = Control=20 not treated, TR1000 TRAIL 1,000 ng/ml, 5 Aza 0.5/1=20 5-Aza-2′-deoxycytidine 0.5/1 μM. Experiments were = performed at least three=20 times. Asterisks outline values which are different from the = respective=20 control (t test, *P < 0.05); n.s. not=20 significant </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec20></A> <DIV class=3DHeading3>Pre-treatment with 5-Aza-2′-deoxycytidine = was not sufficient=20 to sensitise SCG to TRAIL-mediated cell death</DIV> NCH421K_SCG and NCH441_SCG were incubated with 0.5 μM of=20 5-Aza-2′-deoxycytidine for 5 days. After this pre-treatment, = the cells were=20 treated with TRAIL or a combination of TRAIL and = 5-Aza-2′-deoxycytidine for=20 additional 72 h. Apoptotic cell death did not change significantly = in TRAIL=20 treated SCGs. Treatment with 5-Aza-2′-deoxycytidine increased the = fraction of=20 apoptotic cells to 16 =B1 5% in NCH421_SCG and to = 13 =B1 5% in=20 NCH441_SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>d).=20 The combination of TRAIL and 5-Aza-2′-deoxycytidine did not = further enhance=20 apoptotic cell death in SCG, indicating that despite the up-regulation = of=20 caspase-8, 5-Aza-2′-deoxycytidine is not sufficient to sensitise = SCGs to=20 TRAIL-induced cell death. </DIV></DIV> <DIV><A name=3DSec21></A> <HR> <DIV class=3Dheading2>Discussion</DIV> In this study we demonstrated that SCGs are resistant to TRAIL = treatment.=20 This finding is in line with previous reports since it has been already = reported=20 that tumour cells, including primary glioma cells, are resistant to the=20 cytotoxic effects of TRAIL [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR19">19</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR20">20</A></CITE>].=20 In addition to TRAIL resistance, glioblastoma cancer stem cells have = been shown=20 to be insensitive to chemotherapy, e.g., temozolomide, carboplatin, = paclitaxel=20 (Taxol) and etoposide (VP16) [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 Also neural stem and progenitor cells were resistant to CD95-induced = cell death,=20 and upon differentiation they became susceptible for CD95-mediated = apoptosis=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR30">30</A></CITE>].=20 In our study we found that the protein synthesis inhibitor = cycloheximide did=20 not sensitise SCGs to TRAIL-mediated cytotoxicity. In contrast, = TRAIL-induced=20 cell death was efficiently enhanced by cycloheximide in NSCGs. Our = results match=20 the data from the literature because it has been reported that = cycloheximide=20 successfully enhanced TRAIL-mediated apoptosis in malignant glioma cell = lines=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR12">12</A></CITE>].=20 Interestingly, cell lines, e.g., U373 glioma cells, that exhibited = suppressed=20 protein levels of caspase-8 could not be sensitised by cycloheximide to=20 TRAIL-mediated apoptosis [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR12">12</A></CITE>],=20 raising the suggestion that caspase-8 might be silenced in SCGs. In line = with=20 this assumption SCGs exhibited low mRNA levels of caspase-8. In = addition, it has=20 been demonstrated that caspase-8 is frequently lost or silenced in human = glioblastoma [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR2">2</A></CITE>]=20 and in a variety of human cancers [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR15">15</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR16">16</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR36">36</A></CITE>].=20 Furthermore, neural stem and progenitor cells known to be resistant to = death=20 receptor-mediated apoptosis induced by CD95 did not express caspase-8 = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR30">30</A></CITE>].=20 However, exogenous or cytokine-mediated expression of caspase-8 was not=20 sufficient to restore their CD95 sensitivity. These authors also found = that in=20 addition to the absence of caspase-8, primitive neural cells expressed = high=20 levels of the death effector domain containing protein PED (also known = as=20 PEA-15), which localised in the DISC and prevented caspase-8 recruitment = and=20 activation [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR30">30</A></CITE>].=20 In the case of neural stem cells, the absence of caspase-8 and high = expression=20 of PED constitute two levels of protection from apoptosis induced by DRs = and=20 inflammatory cytokines in neural stem and progenitor cells. We also = identified a=20 potential mechanism that led to suppression of caspase-8. We found that = the=20 promoter region of CASP8 in SCGs was methylated as demonstrated = by=20 methylation-specific polymerase chain reaction assay. Treatment with=20 5-Aza-2′-deoxycytidine restored caspase-8 protein in both SCG cell = cultures.=20 Conversely, NSCGs had no methylation of the CASP8 promoter. Our results = match=20 published findings in gliomas demonstrating silenced expression of = caspase-8 due=20 to methylation of the CASP8 promoter [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR2">2</A></CITE>].=20 In neuroblastoma, inactivation of CASP8 by hypermethylation is = the=20 hallmark of a defective apoptosis machinery in advanced disease, = suggesting that=20 CASP8 may act as a tumour suppressor gene in this cancer = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR15">15</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR16">16</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR36">36</A></CITE>].=20 In addition, we demonstrated that in SCGs caspase-8 was also = significantly=20 suppressed on the protein level. In previous reports it has been shown = that=20 tumour cells, including primary glioma cells, that are deficient of or = express=20 only low levels of caspase-8 are completely resistant to TRAIL and that=20 induction of caspase-8 is indispensable for TRAIL-induced apoptosis = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR2">2</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR9">9</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR10">10</A></CITE>].=20 This led us to the conclusion that the suppressed levels of caspase-8 in = SCGs=20 might also explain the high resistance of SCGs to TRAIL-mediated cell = death. We=20 could falsify this hypothesis by restoring caspase-8 expression in both = SCGs by=20 reversing promoter methylation of CASP8 without reversing the = high=20 resistance of these cells towards TRAIL. This contrasts the results of a = previous study in which primary glioma cell lines have been reported to = be=20 resistant to TRAIL stimulation because they expressed low levels of = caspase-8=20 and high levels of the death receptor inhibitor PED/PEA-15 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR7">7</A></CITE>].=20 Treatment with 5-Aza-2′-deoxycytidine resulted in considerable = up-regulation of=20 caspase-8 and sensitisation of primary glioblastoma cells to = TRAIL-induced=20 apoptosis [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR7">7</A></CITE>].=20 Exogenous caspase-8 expression was the main event able to restore TRAIL=20 sensitivity in primary glioblastoma cells. The antitumour activity of=20 5-Aza-2′-deoxycytidine and TRAIL was confirmed in vivo in a mouse = model of=20 glioblastoma multiforme. Evaluation of tumour size, apoptosis and = caspase=20 activation in nude mouse glioblastoma multiforme xenografts showed = dramatic=20 synergy of 5-Aza-2′-deoxycytidine and TRAIL in the treatment of = glioblastoma,=20 whereas the single agents were scarcely effective in terms of reduction = of=20 tumour mass, apoptosis induction and caspase activation. Thus, the = combination=20 of TRAIL and demethylating agents may provide a key tool to overcome=20 glioblastoma resistance to therapeutic treatments. Our results strongly = indicate=20 that in human glioblastoma this combination is insufficient to target = the cancer=20 stem cell fraction. Therefore, TRAIL-based therapies will not fully = eradicate=20 glioblastomas since the cancer stem cell fraction within the tumour is = not=20 efficiently targeted by TRAIL. Hence, treatment strategies are needed = that=20 potently sensitise SCGs to TRAIL-induced cell death. To date, no = treatment=20 strategies specifically sensitising SCGs to TRAIL-mediated cell death = have been=20 described. For primary glioma cell lines such a treatment strategy was = defined=20 and involved the proteasome inhibitor bortezomib [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR21">21</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR22">22</A></CITE>].=20 Bortezomib potently sensitised 13 primary glioma cultures to = TRAIL-mediated=20 apoptosis [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR21">21</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR22">22</A></CITE>].=20 One potential limitation of this study was that they did not demonstrate = that=20 bortezomib and TRAIL also induced apoptosis and synergized in vivo. We also analysed c-FLIP protein, an inhibitor of death = receptor-mediated=20 apoptosis, and detected no difference between SCG and NSCG cells. This = contrasts=20 the results of a recent study in which it was demonstrated that c-FLIP = is=20 significantly up-regulated in CD133+ Jurkat (lymphoma) and CD133+ MCF-7 = (breast=20 carcinoma) cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR40">40</A></CITE>].=20 Down-regulation of FLIP by specific siRNA overcame TRAIL resistance in = CD133+=20 Jurkat and MCF-7 cells, suggesting a major role for c-FLIP in mediating = TRAIL=20 resistance in CD133+ cancer cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR40">40</A></CITE>].=20 However, caspase-8 was not silenced or deleted in CD133+ positive Jurkat = and=20 MCF-7 cells. To identify further factors that possibly convey TRAIL resistance we = employed=20 a proteome array consisting of 30 apoptosis-related proteins. = Interestingly only=20 2 of the 30 proteins were differentially expressed in NSCG and SCG in = both=20 paired cultures, FAS/TNFSF6 and phospho-p53, both of which are not = directly=20 involved in TRAIL-mediated apoptosis. However, in one of the NSCG we = found that=20 TRAIL R2/DR5 was up-regulated. Up-regulation of DR5 has been reported to = confer=20 higher sensitivity to TRAIL [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR1">1</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR29">29</A></CITE>].=20 Recently, it has been demonstrated that the proteasome inhibitor PS-341=20 (bortezomib) up-regulated DR5 expression leading to induction of = apoptosis and=20 enhancement of TRAIL-induced apoptosis despite up-regulation of c-FLIP = and=20 survivin expression in human lung cancer cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR25">25</A></CITE>].=20 Furthermore, it has been reported that the farnesyltransferase inhibitor = R115777=20 up-regulated the expression of DR5 and thereby enhanced TRAIL-mediated = apoptosis=20 in human lung cancer cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR29">29</A></CITE>].=20 Conversely, down-regulation of DR5 by specific siRNA has been shown to = confer=20 TRAIL resistance, suggesting a major role for DR5 in the sensitivity of = cancer=20 cells to TRAIL [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR29">29</A></CITE>].=20 The inhibitor of apoptosis protein (IAP) XIAP, known to inhibit = apoptosis at the=20 level of effector caspase-3/7, was significantly reduced in only one = NSCG. In=20 previous reports XIAP has been linked to confer resistance to TRAIL in = various=20 tumour cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR37">37</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR38">38</A></CITE>].=20 Recently, it has been demonstrated that CD133+ SCGs exhibited higher = XIAP mRNA=20 levels than CD133− glioma cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 For the smallest of the IAPs, survivin, we detected no difference = between SCG=20 and NSCG, although it has been reported that the mRNA level of survivin = in=20 CD133+ positive SCGs was significantly higher than in CD133− = glioma cells=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 Taken together, SCGs were highly resistant towards TRAIL, and this=20 coincidenced with silenced expression of caspase-8, a factor known to be = indispensable for TRAIL-mediated cell death. Reversing methylation of = the=20 caspase-8 promoter led to up-regulation of caspase-8, but failed to = sensitise=20 SCGs to TRAIL-mediated apoptosis. We conclude that whilst treatment with = demethylating agents such as 5-Aza-2′-deoxycytidine sensitises the = bulk tumour=20 cells of glioblastoma, this strategy fails to break the resistance of = CD133+ SCG=20 cells towards TRAIL-mediated apoptosis. As increasing evidence implies = the=20 importance of targeting SCG for an enduring control of glioblastoma, = future=20 research should concentrate on SCG specific sensitisers for = TRAIL-mediated=20 apoptosis. </DIV> <DIV class=3DAcknowledgments><SPAN=20 class=3DAcknowledgmentsHeading>Acknowledgments We= thank=20 Volker Ehemann for performing flow cytometry. 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From: Subject: 10.1007/s00401-009-0494-3 Date: Fri, 6 Mar 2009 09:57:23 +0100 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0000_01C99E41.F2A87940" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0000_01C99E41.F2A87940 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.springerlink.com/content/p536n18xr82w5576/fulltext.html 10.1007/s00401-009-0494-3</TITLE= > <META content=3D"text/html; charset=3DISO-8859-1" = http-equiv=3DContent-Type><LINK=20 rel=3DstyleSheet type=3Dtext/css=20 href=3D"http://www.springerlink.com/config/css/A++_common.css"><LINK=20 rel=3DstyleSheet type=3Dtext/css=20 href=3D"http://www.springerlink.com/config/css/content_A++_SpringerLink.c= ss"> <META name=3DGENERATOR content=3D"MSHTML 8.00.6001.18372"></HEAD> <BODY class=3Dfor-print> <TABLE border=3D1 cellPadding=3D2> <TBODY> <TR class=3Dheader> <TD>Acta Neuropathologica</TD></TR> <TR> <TD>=A9 Springer-Verlag 2009</TD></TR> <TR> <TD>10.1007/s00401-009-0494-3</TD></TR></TBODY></TABLE> <H2 class=3Drubric>Original Paper</H2> <DIV class=3DHeading1><A name=3Dtitle></A>Stem-cell-like glioma cells = are resistant=20 to TRAIL/Apo2L and exhibit down-regulation of caspase-8 by promoter = methylation=20 </DIV> David Capper1,=20 Timo Gaiser1, Christian Hartmann1, 3,=20 Antje Habel1, Wolf Mueller1,=20 Christel Herold-Mende2, = Andreas von Deimling1,=20 3 and Markus David Siegelin1 <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#ContactOfAuthor8"><IMG=20 border=3D0 alt=3D"Contact Information"=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/contact.gif">= </A> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff1></A>(1) </TD> <TD>Department of Neuropathology, = University=20 Hospital Heidelberg, Im Neuenheimer Feld 220, = 69120 Heidelberg,=20 Germany</TD></TR></TBODY></TABLE> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff2></A>(2) </TD> <TD>Department of Neurosurgery, University = Hospital Heidelberg, Im Neuenheimer Feld 400, = 69120 Heidelberg,=20 Germany</TD></TR></TBODY></TABLE> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff3></A>(3) </TD> <TD>Clinical Cooperation Unit = Neuropathology,=20 German Cancer Center (DKFZ) G380, Im Neuenheimer Feld 220,=20 69120 Heidelberg, Germany</TD></TR></TBODY></TABLE> <A name=3DContactOfAuthor8></A> <TABLE class=3DContact> <TBODY> <TR> <TD vAlign=3Dtop><IMG border=3D0 alt=3D"Contact Information"=20 = src=3D"http://www.springerlink.com/content/p536n18xr82w5576/contact.gif">= </TD> = <TD>Markus David Sieg= elin Email:=20 <A=20 = href=3D"mailto:markus.siegelin@med.uni-heidelberg.de">markus.siegelin@med= .uni-heidelberg.de</A></TD></TR></TBODY></TABLE> Received:=20 18 November 2008 Revised:=20 1 February 2009 Accepted:=20 1 February 2009 Published online:=20 12 February 2009 <DIV class=3DAbstract><A name=3DAbs1></A><SPAN=20 class=3DAbstractHeading>Abstract Tumour necrosis = factor=20 (TNF)-related apoptosis-inducing ligand (TRAIL/Apo2L) is a promising = cancer=20 drug. However, many tumours are resistant to TRAIL-based therapies. = Glioma cells=20 with stem cell features (SCG), such as CD133 expression and neurosphere=20 formation, have been recently identified to be more resistant to = cytotoxic drugs=20 than glioma cells lacking stem-cell-like features (NSCGs). Here we = report that=20 SCGs are completely resistant to 100=962,000 ng/ml TRAIL, whereas = NSCGs=20 revealed a moderate sensitivity to TRAIL. We found that SCGs exhibited = only low=20 levels of caspase-8 mRNA and protein, known to be indispensable for=20 TRAIL-induced apoptosis. In addition, we detected hypermethylation of=20 CASP8 promoter in SCGs, whereas NSCGs exhibited a non-methylated=20 CASP8 promoter. Reexpression of caspase-8 by = 5-Aza-2′-deoxycytidine was=20 not sufficient to restore TRAIL sensitivity in SCGs cells, suggesting = that=20 additional factors cause TRAIL resistance in SCGs. Our data suggest that = therapy=20 with TRAIL, either as monotherapy or in combination with demethylating = agents,=20 is not effective in treating glioblastoma because SCGs are not targeted = by such=20 treatment. </DIV> <SPAN=20 class=3DKeywordHeading>Keywords Stem-cell-like glioma=20 cells - Caspase-8 - TRAIL - Apoptosis = - CD133=20 <DIV class=3DKeyword><SPAN=20 class=3DKeywordHeading>Abbreviations <SPAN=20 class=3DTerm>IAP Inhibitor of apoptosis protein - <SPAN=20 class=3DTerm>TRAIL Tumour necrosis factor (TNF)-related=20 apoptosis-inducing ligand - SCG Stem-cell-like=20 glioma cells - NSCG Non-stem-cell-like = glioma cells=20 </DIV> <DIV><A name=3DSec1></A> <HR> <DIV class=3Dheading2>Introduction</DIV> Malignant gliomas are the most common malignant adult primary brain = tumours.=20 Median survival of glioblastoma patients is ~12=9615 months. = Current=20 treatment of malignant glioma usually comprises surgical resection or = diagnostic=20 biopsy followed by adjuvant radiation and chemotherapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR34">34</A></CITE>].=20 Recent studies reported on small sub-populations of tumour cells, = designated=20 as cancer stem cells (CSCs), differing from the majority of malignant = cells=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR32">32</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>].=20 These CSCs are thought to be more resistant to apoptosis, to survive = therapy,=20 and to eventually give rise to recurrent tumours, which usually are = highly=20 resistant to first-line therapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 The reasons for resistance of CSCs to therapy are not well known at = present,=20 although recent research indicates possible molecular determinants = underlying=20 this resistance, e.g., higher expression of BCRP1 and MGMT, as well as = the=20 anti-apoptosis protein and inhibitors of apoptosis protein families = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 Several proteins have been suggested as markers of CSCs, of which CD133 = has=20 received considerable attention because of its relatively high = expression in=20 CSCs of a variety of tumour types, including medulloblastomas [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>],=20 glioblastomas [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>],=20 colorectal [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR28">28</A></CITE>]=20 and prostate cancer [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR5">5</A></CITE>].=20 It has been shown that CD133+ cells, following their separation from = cultured=20 cancer cells or primary tumours, can give rise to glioblastomas in=20 immunocompromised (NOD/SCID) mice, which are phenotypically similar to = primary=20 tumours [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>].=20 It was reported that as many as 107 CD133-cultured = glioblastoma cells=20 were required to form tumours in NOD/SCID mice, whilst as few as = 103=20 CD133+ cells formed tumours in the same model [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR32">32</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR33">33</A></CITE>].=20 Tumour necrosis factor (TNF)-α-related apoptosis-inducing ligand = (TRAIL)=20 belongs to the TNF cytokine family and is capable of inducing apoptosis = in a=20 variety of cancer cells, whilst producing negligible effects on normal = cells=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR11">11</A></CITE>].=20 TRAIL binds to the death receptors DR4/DR5, which subsequently interact = with the=20 adaptor protein FADD and procaspase-8, forming the death-inducing = signalling=20 complex (DISC). Procaspase-8 activation in the DISC leads to cleavage of = procaspase-3 and engagement of the cellular machinery associated with = the type I=20 extrinsic apoptotic pathway [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR4">4</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR27">27</A></CITE>].=20 Activation of the intrinsic, mitochondrial-associated type II apoptotic = pathway=20 is another hallmark of TRAIL-induced cell death because TRAIL, through=20 caspase-8, activates Bid and synergizes with agents that induce = apoptosis=20 exclusively through a type II mechanism [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR6">6</A></CITE>].=20 In type I cells, stimulation of the extrinsic pathway is sufficient for=20 commitment of apoptotic cell death. In type II cells this commitment = requires=20 further signal amplification through the intrinsic pathway [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR23">23</A></CITE>].=20 Caspase-8 resides in the cytosol as a catalytically inactive zymogen and = is=20 present in two isoforms (p55, p53) [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR31">31</A></CITE>].=20 Caspase-8 zymogens are recruited by FADD to the DISC where they are = cleaved=20 through autoproteolysis, releasing the catalytically active p18 and p10 = subunits=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR31">31</A></CITE>].=20 The caspase-8 cleavage can be inhibited by cellular FADD-like=20 interleukin-1β-converting enzyme-inhibitory protein (c-FLIP) = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR17">17</A></CITE>].=20 There are two isoforms of c-FLIP: a short form (c-FLIPS) and = a long=20 form (c-FLIPL). Both c-FLIPS and = c-FLIPL are=20 recruited to the DISC where c-FLIPL is cleaved into a p43=20 intermediate form, which, together with c-FLIPS, remains in = the DISC=20 where they inhibit caspase-8 cleavage [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR39">39</A></CITE>].=20 Epigenetic alterations in DNA have been shown to be important in the = genesis=20 and progression of tumours, especially the methylation-mediated = silencing of=20 tumour suppressor genes [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR18">18</A></CITE>].=20 In cancer cells, aberrant methylation of CpG islands located in the = promoter=20 regions of genes implicated in apoptosis, migration, or DNA repair is = frequently=20 associated with transcriptional silencing and gene repression [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR18">18</A></CITE>].=20 The former alterations of the =91epigenome=92 also contribute to = defining the=20 biological behaviour of the tumour and might modulate the response of = tumour=20 cells to anticancer therapies [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR18">18</A></CITE>].=20 In the present study we found that CD133+ stem-cell-like glioblastoma = cells=20 (SCG) were completely resistant towards treatment with TRAIL and = exhibited only=20 low levels of caspase-8 mRNA and protein compared to CD133− = non-stem-cell-like=20 glioblastoma cells (NSCG). The up-regulation of caspase-8 by=20 5-Aza-2′-deoxycytidine treatment was not sufficient to sensitise = SCG to TRAIL,=20 indicating the existence of further factors mediating TRAIL resistance = in SCG.=20 </DIV> <DIV><A name=3DSec2></A> <HR> <DIV class=3Dheading2>Materials and methods</DIV> <DIV><A name=3DSec3></A> <DIV class=3DHeading3>Cell culture and reagents</DIV> Two paired cultures, NCH421K_SCG/NCH421K_NSCG and = NCH441_SCG/NCH441_NSCG,=20 were derived from surgical glioblastoma samples. The generation and=20 characteristics of these cell lines are described in detail elsewhere. = In brief,=20 glioma cell cultures derived from surgically removed tumour tissue were = FACS=20 sorted (see below) for CD133 expression and separated in CD133+ (SCG) = and CD133−=20 (NSCG) tumour cell fractions and were either established in foetal calf=20 serum-containing medium (NSCG) or a medium optimised for the growth of = normal=20 neural stem cells containing bFGF, EGF and the serum-free supplement = BIT9500=20 (SCG). SCG cells displayed stem cell typical growth as neurospheres, = increased=20 expression of CD133 and nestin, and were highly tumorigenic after = implantation=20 in NOD/SCID mice; 103 SCGs were sufficient to form tumours in = nude=20 mice, whereas 106 NSCG cells were necessary to achieve tumour = growth=20 in vivo [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR35">35</A></CITE>].=20 Common origin of the paired cell lines was confirmed by microsatellite = analysis=20 targeting polymorphisms on chromosomes 1 and 19 as previously described=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR13">13</A></CITE>].=20 NCH421K_NSCG and NCH441_NSCG were cultured in DMEM Glutamax-I = 4,500 g/l=20 glucose (Invitrogen, Karlsruhe, Germany) with 10% FBS and 1%=20 penicillin/streptomycin (Invitrogen, Karlsruhe) and were incubated at = 37=B0C in a=20 humidified atmosphere containing 10% carbon dioxide. NCH421_SCG and = NCH441_SCG=20 were incubated with serum-free neural stem cell medium containing = DMEM/F12=20 (Gibco), recombinant human epidermal growth factor (rhEGF, = 20 ng/ml; Sigma,=20 USA), basic fibroblast growth factor (bFGF, 20 ng/ml; Upstate, = USA), BIT:=20 bovine serum albumin, transferrin, insulin and 100 U/ml penicillin = G=20 (Provitro, Germany) and 100 μg/ml streptomycin (Gibco, USA). = Recombinant=20 human TRAIL/Apo2L was purchased from Peprotech (Rocky Hill, NY). = Cycloheximide=20 was obtained from Axxora (Loerrach, Germany), and cells were treated = with the=20 indicated amounts of cycloheximide as individually indicated. To exclude = the=20 effects of culture conditions on TRAIL resistance of SCG cells, SCG cell = lines=20 were additionally cultured for 48 h in DMEM Glutamax-I = 4,500 g/l=20 glucose with 10% FBS and 1% penicillin/streptomycin prior to TRAIL = treatment in=20 selected experiments without observable differences to culture = conditions in=20 neural stem cell medium. </DIV> <DIV><A name=3DSec4></A> <DIV class=3DHeading3>Caspase-3 activity assay</DIV> Cells were collected and analysed by Caspase-3 Colorimetric Assay Kit = (R&D Systems, Minneapolis, MN). Cells were lysed to collect their=20 intracellular contents. The cell lysate was tested for protease activity = by the=20 addition of a caspase-specific peptide that was conjugated to the colour = reporter molecule p-nitroanaline (pNA). The cleavage of the = peptide by=20 the caspase releases the chromophore pNA, which was quantitated=20 spectrophotometrically at a wavelength of 405 nm. The level of = caspase=20 enzymatic activity in the cell lysate was directly proportional to the = colour=20 reaction. </DIV> <DIV><A name=3DSec5></A> <DIV class=3DHeading3>Cell viability</DIV> Cells were seeded into 96-well plates at a density of=20 2 =D7 104 cells/well in 100-μl tissue = culture medium in=20 triplicate. After 24-h incubation allowing cells to adhere, cells were = treated=20 for 24 h either with cycloheximide or TRAIL or with combinations of = both,=20 as described in individual experiments. Cell viability was determined by = the=20 colorimetric = dimethylthiazolyl-carboxymethoxyphenyl-sulphophenyl-tetrazolium=20 (MTS) assay. In this assay, the quantity of formazan product formed is = directly=20 proportional to the number of viable cells in the cultures. After = 4 h in=20 culture the cell viability was determined by measuring the absorbance at = 490 nm using a 550 Bio-Rad plate-reader (Bio-Rad, Hertfordshire, = UK). The=20 relative percentage of survival was calculated by dividing the = absorbance of=20 treated cells by that of the control in each experiment. IC50 = values=20 (concentrations that led to a reduction of 50% cellular viability) were=20 calculated. </DIV> <DIV><A name=3DSec6></A> <DIV class=3DHeading3>Annexin V/PI staining</DIV> Apoptotic cells were assessed by annexin V staining using Annexin = V-FITC=20 apoptosis detection kit (BD Bioscience) following the manufacturer=92s=20 instructions and analysed by flow cytometry using Cellquest (Becton = Dickinson).=20 In the annexin V assay, the cells staining only for Annexin V-FITC = (early=20 apoptosis) are located in the lower right quadrant, whereas cells = staining for=20 both annexin V and propidiumiodide (PI) (late apoptosis) are located in = the=20 upper left quadrant. The percent positive cells for PI staining only in = the top=20 left quadrant represent the necrotic population. </DIV> <DIV><A name=3DSec7></A> <DIV class=3DHeading3>Flow cytometric analysis of protein expression and = cell=20 sorting</DIV> The level of expression of CD133 protein was assessed by flow = cytometry. In=20 brief, cells were harvested. They were then incubated with anti-CD133/2 = IgG=20 (clone 293C3; Miltenyi Biotec, Bergisch-Gladbach, Germany) followed by=20 incubation with secondary FITC- or PE-conjugated IgG (R&D Systems,=20 Wiesbaden, Germany). The cells were then sorted by flow cytometry = (FACSCalibur,=20 Becton Dickinson) for populations with high and low fluorescence. = </DIV> <DIV><A name=3DSec8></A> <DIV class=3DHeading3>Real-time reverse transcription PCR</DIV> Total RNA was isolated using the RNA Nucleospin RNA-II kit = (Macherey-Nagel,=20 D=FCren, Germany) according to the manufacturer=92s instructions. The = total RNA=20 concentration was assessed using a Nano-Drop photometer (Peqlab = Biotechnologie,=20 Erlangen, Germany). For cDNA synthesis 1 μg of total RNA was = used for=20 reverse transcription (random primer) using RevertAidTM H Minus MuLV = Polymerase=20 (1 U/μl), according to the manufacturer=92s instructions (MBI = Fermentas, St.=20 Leon-Rot, Germany). The following primer sequences were used: caspase-8 = forward,=20 5′CAG CAG CCT TGA AGG AAG TC3′; caspase-8 reverse, = 5′CGA GAT TGT CAT TAC CCC=20 ACA3′; c-FLIP forward, 5′CTC ACC GTC CCT GTA CCT3′; = c-FLIP reverse, 5′CAG GAG=20 TGG GCG TTT TCT3′; GAPDH forward, 5′TGC CTC TTT AGT TGT CAT = GCA G3′; GAPDH=20 reverse, 5′CCC GTT CAG CTC AGG GAT GA3′. All real-time PCR = reactions were=20 performed in a 25-μl mixture containing 1/20 volume of cDNA = preparation=20 (1 μl), 1=D7 SYBR Green buffer (PE Applied Biosystems, Foster = City, CA),=20 4 mM MgCl2, 0.2 μM of each primers (BA67 and = BA68),=20 0.2 mM dNTPs mix and 0.025 Unit of AmpliTaq Gold=AE = thermostable=20 DNA polymerase (Applied Biosystems, Foster City, CA). The PCR was run on = an ABI=20 PRISM Detection Instrument 7400. After 40 cycles, data reduction was = performed=20 with Sequence Detection System Software (Applied Biosystems Inc., Foster = City,=20 CA). For data analysis, threshold cycles (C T) for = GAPDH=20 (reference) and c-FLIP or caspase-8 (sample) were determined in = triplicates. We=20 chose cDNA of NCG421K_NSCG and NCH441_NSCG as calibrator cells (100%) = and=20 determined the relative change (rI) in copy numbers according to the = formula=20 rI =3D 2 exp[(C T c-FLIP or caspase-8=20 NSCG − C TGAPDH = NSCG) − (C=20 T c-FLIP or caspase-8 SCG − C = TGAPDH=20 SCG)]. </DIV> <DIV><A name=3DSec9></A> <DIV class=3DHeading3>Methylation-specific polymerase chain reaction = assay</DIV> DNA methylation patterns in the CpG islands of CASP8 were = determined=20 by chemical modification of unmethylated, but not the methylated, = cytosines to=20 uracil and subsequent polymerase chain reaction using primers specific = for=20 either methylated or the modified unmethylated DNA [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR8">8</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR14">14</A></CITE>].=20 One microgram of DNA was bisulphite treated and purified with the = EpiTect=20 Bisulfite Kit (QIAGEN Inc., Valencia, CA) following the manufacturer=92s = instructions. Primer sequences for methylation analysis are as described = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR26">26</A></CITE>]=20 (unmethylated sense: 5′-GTT GGT TTT ATT TAG TTT GGT-3′; = unmethylated antisense:=20 5′-CCC TAT CAA TAA CAA ATA ATA TAC-3′; methylated sense: = 5′-GTT GGT TTT ATT TAG=20 TTC GGC-3′; methylated antisense: 5′-CCC TAT CGA TAA CAA ATA = ATA TAC-3′).=20 Placental DNA treated in vitro with Sss I methyltransferase (New England = BioLabs, Beverly, MA) was used as positive control for methylated = alleles, and=20 DNA from normal lymphocytes was used as negative control for methylated = alleles.=20 Controls without DNA were performed for each set of polymerase chain = reaction.=20 Ten microlitres of each polymerase chain reaction was directly loaded = onto=20 non-denaturing 2% agarose gels, stained with ethidium bromide and = visualised=20 under UV illumination. </DIV> <DIV><A name=3DSec10></A> <DIV class=3DHeading3>Western blot</DIV> Twenty micrograms of protein diluted in NuPAGE-sample buffer and = reducing=20 reagent (Invitrogen, Carlsbad, Germany) were denatured at 95=B0C for = 5 min=20 and electrophoretically separated on ready-to-use 4=9612% sodium dodecyl = sulphate=96polyacrylamide gel electrophoresis (SDS=96PAGE) (Invitrogen, = Carlsbad).=20 Proteins were blotted onto nitrocellulose membranes at=20 1.5 mA/cm2 for 1.5 h (Invitrogen, Carlsbad). After = blocking=20 in 0.5 M Tris=96Base, pH 7.4, 5% milk powder, 1.5 M NaCl and = 0.05%=20 Tween, the membranes were incubated with mouse anti-human caspase-8 = antibody=20 diluted 1:1,000 (clone 1C12; CST, Danvers, MA), rabbit anti-human PARP = (clone=20 46D11; CST, Danvers), rabbit anti-human c-FLIP (R&D Systems, = Minneapolis,=20 MN), rabbit anti-cleaved caspase-3 (CST, Danvers) and rabbit anti-human=20 caspase-7 (clone Asp198; CST Inc., Danvers) overnight at 4=B0C. Staining = with=20 secondary horseradish peroxidase conjugated anti-rabbit or anti-mouse = antibodies=20 at dilutions of 1:10,000 or 1:2,000, respectively (Amersham Biosciences, = Buckinghamshire, UK) was followed by immunodetection with Western = Blotting=20 Detection System ECL (Amersham Biosciences, Buckinghamshire). Protein = signals=20 were analysed semiquantitatively, using a computer-assisted image = analysis=20 system and the NIH gel analysis software (<A=20 href=3D"http://www.rsb.info.nih.gov/nihimage/download.html">http://www.rs= b.info.nih.gov/nihimage/download.html</A>).=20 </DIV> <DIV><A name=3DSec11></A> <DIV class=3DHeading3>Human Apoptosis Array Kit/Proteome = Profiler=99</DIV> For analysing the expression profiles of apoptosis-related proteins, = we used=20 the Human Apoptosis Array Kit (R&D Systems Ltd., Abingdon, UK). This = assay=20 consists of 30 nitrocellulose membrane spotted antibodies specific for=20 apoptosis-related proteins. Cell samples (1 =D7 107 = cells)=20 were harvested, and 300 μg of protein was mixed with = 15 μl of=20 biotinylated detection antibodies. After pre-treatment the cocktail was=20 incubated overnight at 4=B0C on a rocking platform. Following a wash = step to=20 remove unbound material, streptavidin=96horseradish and chemiluminescent = detection=20 reagents were added sequentially. Chemiluminescence at each spot is = proportional=20 to the amount of bound substrate. All experiments were repeated three = times. The=20 signals on X-ray film were quantified by scanning on a transmission-mode = scanner=20 and analysing the array image file using ImageJ analysis software (<A=20 href=3D"http://rsbweb.nih.gov/ij/">http://rsbweb.nih.gov/ij/</A>). In = detail, the=20 following procedure was carried out: The averaged background signal was subtracted and then the arrays = were=20 calibrated based on the signal strength of the positive controls. The = average=20 signal (pixel density) of the pair of duplicated spots representing each = apoptosis protein was determined. The corresponding signals on different = arrays=20 were compared and the relative change in apoptosis proteins was = determined.=20 </DIV></DIV> <DIV><A name=3DSec12></A> <HR> <DIV class=3Dheading2>Results</DIV> <DIV><A name=3DSec13></A> <DIV class=3DHeading3>SCGs express CD133</DIV> <DIV class=3DPara> <DIV>NCH421K_SCG, NCH421K_NSCG, NCH441_SCG and NCH441_NSCG were analysed = for the=20 expression of CD133 by flow cytometry. Two representative flow cytometry = charts=20 are shown (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig1">1</A>a,=20 b). In stem-cell-like glioma cell culture, 91% of NCH421K_SCG and 92% = NCH441_SCG=20 stained positive for CD133 (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig1">1</A>c).=20 In contrast, NCH421K_NSCG and NCH441_NSCG cultures contained only 1 and = 2% CD133=20 positive cells, respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig1">1</A>c).=20 <DIV class=3DFigure><A name=3DFig1></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig1_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig1_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 1 Rate of = CD133 in SCGs=20 and NSCGs. Flow cytometry of NCH421K_SCG (a) and NCH441_SCG = (b)=20 stained with CD133 antibody. M2 represents cells positive, whereas M1 = shows=20 cells negative for CD133. c Diagram of CD133 rate in NCH421K_SCG, = NCH441_SCG, NCH421K_NSCG and NCH441_NSCG determined by flow cytometry = </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec14></A> <DIV class=3DHeading3>SCGs are resistant towards TRAIL treatment</DIV> <DIV class=3DPara> <DIV>SCGs were resistant to TRAIL treatment (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>).=20 Treatment with 100, 250, 500, 1,000 and 2,000 ng/ml TRAIL for = 24 h=20 alone did not have a significant effect on cell death in NCH441_SCG=20 (IC50 > 2,000 ng/ml) and NCH421K_SCG=20 (IC50 > 2,000 ng/ml) (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>a).=20 Even prolonged treatment with TRAIL for up to 7 days did not induce = cell=20 death in SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>b).=20 In contrast, NSCGs were moderately sensitive to TRAIL. Treatment with = different=20 concentrations of TRAIL for 24 h had a significant effect on cell = death in=20 NCH441_NSCG and NCH421_NSCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>a).=20 The IC50s were calculated for NCH441_NSCG=20 (220 =B1 40 ng/ml) and NCH421K_NSCG=20 (177 =B1 52 ng/ml).=20 <DIV class=3DFigure><A name=3DFig2></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig2_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig2_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 2 MTT = viability assay=20 and Western blot analysis after treatment with TRAIL, cycloheximide or = the=20 combination of both after 24 h of treatment. a MTT assay in=20 NCH421K_SCG, NCH441_SCG, NCH421K_NSCG and NCH441_NSCG cells upon = treatment with=20 TRAIL for 24 h. b MTT assay in NCH421K_SCG and NCH441_SCG = upon=20 treatment with 1,000 ng/ml TRAIL for up to 7 days. c = MTT assay=20 in NCH421K_SCG, NCH441_SCG, NCH421K_NSCG and NCH441_NSCG cells upon = treatment=20 with TRAIL, cycloheximide or the combination of both for 24 h. = d=20 Western blot showing cleaved caspase-7 and PARP in NCH421K_SCG, = NCH441_SCG,=20 NCH421K_NSCG and NCH441_NSCG untreated or treated with 500 ng/ml = TRAIL for=20 8 h. e Caspase-3 activity in NCH421K_SCG, NCH441_SCG, = NCH421K_NSCG=20 and NCH441_NSCG upon treatment with TRAIL. Control not treated,=20 TR100/250/500/1000/2000 TRAIL 100/250/500/1,000/2,000 ng/ml, = 0d/1d/2d/5d/7d day 0/1/2/5/7 after start of treatment with TRAIL, = CHX20 cycloheximide 20 μg/ml. Experiments were = performed at least=20 three times. Asterisks outline values that are different from the = respective control (t test, *P < 0.05); = n.s.=20 not significant </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec15></A> <DIV class=3DHeading3>Inhibition of protein synthesis by cycloheximide = does not=20 sensitise SCGs to TRAIL-mediated cytotoxicity, but enhances = TRAIL-induced=20 cytotoxicity in NSCGs </DIV> After 24 h treatment with cycloheximide 20 μg/ml cell = death=20 increased significantly in NCH441_SCG (27 =B1 3%) and = NCH421K_SCG=20 (36 =B1 3%), respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Combining 20 μg/ml cycloheximide with 1,000 ng/ml of = TRAIL did not=20 increase cell death significantly in NCH441_SCG (30 =B1 3%) = and=20 NCH421K_SCG (44 =B1 3%) compared to single treatment with = cycloheximide=20 (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Thus, the protein synthesis inhibitor was not capable of sensitising = SCGs to=20 TRAIL-mediated cytotoxicity. Single treatment with 20 μg/ml cycloheximide increased cell = death in=20 NCH441_NSCG (18 =B1 3%) and NCH421K_NSCG = (17 =B1 3%),=20 respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Combining 20 μg/ml cycloheximide with 100 ng/ml of = TRAIL increased=20 cell death significantly in NCH441_NSCG (65 =B1 3%) and = NCH421K_NSCG=20 (74 =B1 3%) compared to single treatment with cycloheximide or = TRAIL-only treated cells (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>c).=20 Thus, in NSCGs the protein synthesis inhibitor further enhanced the = cytotoxic=20 effect of TRAIL. </DIV> <DIV><A name=3DSec16></A> <DIV class=3DHeading3>TRAIL mediates activation (cleavage) of = effector-caspases 3,=20 7 and poly-ADP-ribose-polymerase in NSCGs</DIV> We employed Western blotting to elucidate the proteolytic mechanism = in=20 TRAIL-induced apoptosis (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>d).=20 NCH441_SCG, NCH421K_SCG, NCH441_NSCG and NCH421K_NSCG cells were treated = with=20 TRAIL for 12 h, and the activation/cleavage of caspase-7 and=20 poly-ADP-ribose-polymerase (PARP) were examined. Exposure of cells to=20 500 ng/ml TRAIL alone yielded undetectable signals of cleaved = fragment of=20 PARP sized 89 kDa and active cleaved caspase-7 sized 20 kDa in = NCH441_SCG and NCH421K_SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>d).=20 Treatment with 500 ng/ml TRAIL led to a significant increase in = cleaved=20 fragment of PARP and active cleaved caspase-7/3 in NCH441_NSCG and = NCH421K_NSCG=20 (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>d).=20 Upon treatment with 500 or 1,000 ng/ml TRAIL SCG did not exhibit an = increase in caspase-3 activity, whereas NSCG showed a significant = increase in=20 caspase-3 activity compared to the controls (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig2">2</A>e).=20 </DIV> <DIV><A name=3DSec17></A> <DIV class=3DHeading3>Human Apoptosis Array Kit/Proteome = Profiler=99</DIV> <DIV class=3DPara> <DIV>To identify the resistance mechanism leading to TRAIL resistance in = SCGs we=20 analysed the expression of 30 apoptosis-related proteins that have = already been=20 shown to be associated with TRAIL resistance (Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>a;=20 Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig3">3</A>a,=20 b). Concentrations of the following proteins were significantly = different in SCG=20 and NSCG: Bad, cleaved caspase-3 Fas/TNFSF6, TRAIL R2/DR5, phospho-p53 = (S392),=20 XIAP, FADD, HSP60 and p21/CIP1/CDNK1A (Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>a,=20 b). DR5 and FADD were significantly increased in NCH441_NSCG compared to = NCH441_SCG, indicating a higher susceptibility of NCH441_NSCG to = TRAIL-mediated=20 apoptosis. However, only 2 of the 30 analysed proteins were = differentially=20 regulated in both cell line pairs. We detected a significant = up-regulation of=20 Fas/TNFSF6 and a down-regulation of phospho-p53 (S392) in NSCGs = (Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>b,=20 c).<A name=3DTab1></A> <DIV class=3DCapt>Table 1 Human = Apoptosis=20 Array Kit/Proteome Profiler=99 </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH colSpan=3D4 align=3Dleft> (a)</TH></TR> <TR class=3Dheader> <TH align=3Dleft> Coordinate</TH> <TH align=3Dleft> Target/control</TH> <TH align=3Dleft> Coordinate</TH> <TH align=3Dleft> Target/control</TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> A1, A2</TD> <TD align=3Dleft> Positive control</TD> <TD align=3Dleft> C13, C14</TD> <TD align=3Dleft> HO-2/HMOX2</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> A23, A24</TD> <TD align=3Dleft> Positive control</TD> <TD align=3Dleft> C15, C16</TD> <TD align=3Dleft> HSP27</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B1, B2</TD> <TD align=3Dleft> Bad</TD> <TD align=3Dleft> C17, C18</TD> <TD align=3Dleft> HSP60</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B3, B4</TD> <TD align=3Dleft> Bax</TD> <TD align=3Dleft> C19, C20</TD> <TD align=3Dleft> HSP70</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B5, B6</TD> <TD align=3Dleft> Bcl-2</TD> <TD align=3Dleft> C21, C22</TD> <TD align=3Dleft> HTRA2/Omi</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B7, B8</TD> <TD align=3Dleft> Bcl-x</TD> <TD align=3Dleft> C23, C24</TD> <TD align=3Dleft> Livin</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B9, B10</TD> <TD align=3Dleft> Procaspase-3</TD> <TD align=3Dleft> D1, D2</TD> <TD align=3Dleft> PON2</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B11, B12</TD> <TD align=3Dleft> Cleaved caspase-3</TD> <TD align=3Dleft> D3, D4</TD> <TD align=3Dleft> p21/CIP1/CDNK1A</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B13, B14</TD> <TD align=3Dleft> Catalase</TD> <TD align=3Dleft> D5, D6</TD> <TD align=3Dleft> p27/Kip1</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B15, B16</TD> <TD align=3Dleft> cIAP-1</TD> <TD align=3Dleft> D7, D8</TD> <TD align=3Dleft> Phospho-p53 (S15)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B17, B18</TD> <TD align=3Dleft> cIAP-2</TD> <TD align=3Dleft> D9, D10</TD> <TD align=3Dleft> Phospho-p53 (S46)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B19, B20</TD> <TD align=3Dleft> Claspin</TD> <TD align=3Dleft> D11, D12</TD> <TD align=3Dleft> Phospho-p53 (S392)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B21, B22</TD> <TD align=3Dleft> Clusterin</TD> <TD align=3Dleft> D13, D14</TD> <TD align=3Dleft> Phospho-Rad17 (S635)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> B23, B24</TD> <TD align=3Dleft> Cytochrome c </TD> <TD align=3Dleft> D15, D16</TD> <TD align=3Dleft> SMAC/Diablo</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C1, C2</TD> <TD align=3Dleft> TRAIL R1/DR4</TD> <TD align=3Dleft> D17, D18</TD> <TD align=3Dleft> Survivin</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C3, C4</TD> <TD align=3Dleft> TRAIL R2/DR5</TD> <TD align=3Dleft> D19, D20</TD> <TD align=3Dleft> TNF RI/TNFRSF1A</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C5, C6</TD> <TD align=3Dleft> FADD</TD> <TD align=3Dleft> D21, D22</TD> <TD align=3Dleft> XIAP</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C7, C8</TD> <TD align=3Dleft> Fas/TNFSF6</TD> <TD align=3Dleft> D23, D24</TD> <TD align=3Dleft> PBS (negative control)</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C9, C10</TD> <TD align=3Dleft> HIF-1</TD> <TD align=3Dleft> E1, E2</TD> <TD align=3Dleft> Positive control</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> C11, C12</TD> <TD align=3Dleft> HO1/HMOX1/HSP32</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD></TR></TBODY></TABLE> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dchar char=3D"."> <COL align=3Dchar char=3D"=B1"> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH colSpan=3D4 align=3Dleft> (b)</TH></TR> <TR class=3Dheader> <TH align=3Dleft> </TH> <TH align=3Dleft char=3D"."> NCH421K_SCG</TH> <TH align=3Dleft char=3D"=B1"> NCH421K_NSCG</TH> <TH align=3Dleft> </TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> Bad </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.56 =B1 0.15 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bax</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.2 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-x</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.0 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Procaspase-3</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.26</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cleaved caspase-3 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 3.43 =B1 0.24 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Catalase</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.10</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Claspin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.15 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Clusterin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cytochrome c </TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R1/DR4</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.07 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R2/DR5</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.10</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> FADD</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Fas/TNFSF6 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 10.34 =B1 0.22 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HIF-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.16</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO1/HMOX1/HSP32</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO-2/HMOX2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.11 =B1 0.24</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP27</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.55 =B1 0.23</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP60</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.23 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP70</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.35 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HTRA2/Omi</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Livin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.13 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> PON2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.22 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p21/CIP1/CDNK1A</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.35 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p27/Kip1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.91 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S15)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.94 =B1 0.17</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S46)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S392) </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.55 =B1 0.21 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-Rad17 (S635)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.2 =B1 0.29</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> SMAC/Diablo</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.22</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Survivin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.27</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TNF RI/TNFRSF1A</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.28</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> XIAP </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.45 =B1 0.26 </TD> <TD align=3Dleft> </TD></TR></TBODY></TABLE> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dchar char=3D"."> <COL align=3Dchar char=3D"=B1"> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH colSpan=3D4 align=3Dleft> (c)</TH></TR> <TR class=3Dheader> <TH align=3Dleft> </TH> <TH align=3Dleft char=3D"."> NCH441_SCG</TH> <TH align=3Dleft char=3D"=B1"> NCH441_NSCG</TH> <TH align=3Dleft> </TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> Bad</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bax</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.28 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.15 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Bcl-x</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.0 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Procaspase-3</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.26</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cleaved caspase-3</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.24</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Catalase</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.10</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.11</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> cIAP-2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.13</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Claspin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.15 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Clusterin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.12 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Cytochrome c </TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.15</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R1/DR4</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.23 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TRAIL R2/DR5 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 1.88 =B1 0.15 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> FADD </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 1.82 =B1 0.18 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Fas/TNFSF6 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 16.12 =B1 0.22 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HIF-1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.16</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO1/HMOX1/HSP32</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.18</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HO-2/HMOX2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.11 =B1 0.24</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP27</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.55 =B1 0.23</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP60 </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.56 =B1 0.13 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HSP70</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.35 =B1 0.19</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> HTRA2/Omi</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.09 =B1 0.28</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Livin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.13 =B1 0.23</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> PON2</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.22 =B1 0.25</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p21/CIP1/CDNK1A </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 8.56 =B1 0.54 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> p27/Kip1</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.91 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S15)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.94 =B1 0.17</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S46)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.95 =B1 0.12</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-p53 (S392) </TD> <TD align=3Dchar char=3D"."> 1 </TD> <TD align=3Dchar char=3D"=B1"> 0.18 =B1 0.23 </TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Phospho-Rad17 (S635)</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.2 =B1 0.29</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> SMAC/Diablo</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 0.96 =B1 0.22</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> Survivin</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.1 =B1 0.27</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> TNF RI/TNFRSF1A</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.14 =B1 0.28</TD> <TD align=3Dleft> </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> XIAP</TD> <TD align=3Dchar char=3D"."> 1</TD> <TD align=3Dchar char=3D"=B1"> 1.13 =B1 0.26</TD> <TD align=3Dleft> </TD></TR></TBODY></TABLE> <DIV class=3DCapt> <DIV class=3DCaptCont> <DIV>a: Identification of the 30 apoptosis-related proteins on the = protein=20 array. b, c: Expression level of 30 apoptosis-related proteins in = NCH421K_SCG=20 (b), NCH441_SCG (c), NCH421_NSCG (b) and NCH441_NSCG (b). Protein levels = of SCG=20 were normalised to 1, and results are given as relative values with=20 corresponding standard deviations. Proteins and values in bold were=20 significantly different. Arrays and analysis were performed thrice=20 </DIV></DIV></DIV> <DIV class=3DFigure><A name=3DFig3></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig3_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig3_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 3 Analysis = of 30=20 apoptosis-related proteins in SCGs and NSCGs. a Stem-cell-like=20 (NCH421K_SCG) and non-stem-cell-like glioblastoma cells (NCH421K_NSCG) = were=20 harvested and incubated with nitrocellulose membranes containing 30=20 apoptosis-related proteins. b Stem-cell-like (NCH441_SCG) and=20 non-stem-cell-like glioblastoma cells (NCH441_NSCG) were harvested and = incubated=20 with nitrocellulose membranes containing 30 apoptosis-related proteins. = Proteins=20 with a significantly different expression are marked and outlined in the = figure.=20 For results of densitometry, refer to Table <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Tab1">1</A>=20 </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec18></A> <DIV class=3DHeading3>Caspase-8 expression is suppressed in SCGs</DIV> Since the human apoptosis array kit does not include c-FLIP and = caspase-8, we=20 analysed the expression of c-FLIP and caspase-8 by real-time reverse=20 transcription PCR and Western blot. <DIV class=3DPara> <DIV>Significantly different levels of c-FLIP mRNA or protein were not = observed=20 between SCGs and NSCGs (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig4">4</A>a=96c).=20 <DIV class=3DFigure><A name=3DFig4></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig4_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig4_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 4 Expression analysis=20 of c-FLIP and caspase-8 mRNA and protein level. a, b mRNA = levels=20 of c-FLIP and caspase-8 were analysed in NCH421K_SCG, NCH421K_NSCG, = NCH441_SCG=20 and NCH441_NSCG. The levels of c-FLIP and caspase-8 were normalised to 1 = in=20 non-stem-cell-like glioma cells. Experiments were repeated thrice. = c=20 Western blot analysis of c-FLIP and caspase-8 in NCH421K_SCG, = NCH421_NSCG,=20 NCH441_SCG and NCH441_NSCG. Actin served as protein loading control = </DIV> <HR> </DIV></DIV> Both SCG cell lines demonstrated significantly reduced levels of = caspase-8=20 compared to NSCGs. In NCH421K_SCG the relative mRNA level of caspase-8 = was=20 76 =B1 3% (P < 0.01) lower than in = NCH421K_NSCG, and=20 in NCH441_SCG it was 98 =B1 5% (P < 0.01) = lower than=20 in NCH441_NSCG, respectively (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig4">4</A>a,=20 b). These results were confirmed on the protein level. The protein = levels of=20 caspase-8 were significantly reduced in both SCGs, NC441_SCG and = NCH421K_SCG,=20 compared to the NSCG cells (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig4">4</A>c).=20 </DIV> <DIV><A name=3DSec19></A> <DIV class=3DHeading3>Hypermethylation of the CASP8 promoter = region in SCGs=20 </DIV> <DIV class=3DPara> <DIV>Caspase-8 was suppressed at the level of transcription. Therefore,=20 silencing of caspase-8 in SCG might be due to methylation of the = CASP8=20 promoter region. We employed methylation-specific polymerase chain = reaction to=20 assess the methylation status of CASP8 as described in the = materials and=20 methods section. Both SCGs, NCH421K_SCG and NCH441_SCG, demonstrated a=20 methylated CASP8 promoter (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>a).=20 In contrast, NSCGs cells exhibited no methylation of the CASP8 = promoter=20 in both cell lines NCH421K_NSCG and NCH441_NSCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>a).=20 Importantly, treatment with 5-Aza-2′-deoxycytidine resulted in a = significant=20 up-regulation of caspase-8 on both mRNA and protein levels in NCH421_SCG = and=20 NCH441_SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>b,=20 c), suggesting that hypermethylation of CASP8 promoter is = responsible for=20 suppression of caspase-8 levels in SCG cells.=20 <DIV class=3DFigure><A name=3DFig5></A><IMG=20 alt=3DMediaObjects/401_2009_494_Fig5_HTML.gif=20 src=3D"http://www.springerlink.com/content/p536n18xr82w5576/MediaObjects/= 401_2009_494_Fig5_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 5 Analysis = of CpG=20 island promoter methylation status in SCGs and NSCGs by the = methylation-specific=20 polymerase chain reaction assay. a The presence of a visible = polymerase=20 chain reaction product in those lanes marked U indicates the = presence of=20 an unmethylated CASP8 promoter; the presence of product in those = lanes=20 marked M indicates the presence of a methylated CASP8 = promoter.=20 Water (H2O) was used as negative polymerase chain reaction = control.=20 In vitro methylated placental DNA (IVD) was used as positive control for = methylated genes. DNA from normal lymphocytes (NLs) was used as negative = control=20 for methylated alleles. b mRNA levels of caspase-8 NCH421_SCG and = NCH441_SCG after treatment with different concentrations (μM) of=20 5-Aza-2′-deoxycytidine (5 Aza) for 120 h. c Protein = levels of=20 caspase-8 after treatment with different concentrations (μM) of=20 5-Aza-2′-deoxycytidine (5 Aza) for 120 h. LN229 is an = established cell line=20 expressing considerable levels of caspase-8. d Annexin V/PI flow=20 cytometric based assay in NCH421K_SCG, NCH441_SCG, upon treatment with = TRAIL,=20 5-Aza-2′-deoxycytidine or the combination of both for 72 h. = Control=20 not treated, TR1000 TRAIL 1,000 ng/ml, 5 Aza 0.5/1=20 5-Aza-2′-deoxycytidine 0.5/1 μM. Experiments were = performed at least three=20 times. Asterisks outline values which are different from the = respective=20 control (t test, *P < 0.05); n.s. not=20 significant </DIV> <HR> </DIV></DIV></DIV> <DIV><A name=3DSec20></A> <DIV class=3DHeading3>Pre-treatment with 5-Aza-2′-deoxycytidine = was not sufficient=20 to sensitise SCG to TRAIL-mediated cell death</DIV> NCH421K_SCG and NCH441_SCG were incubated with 0.5 μM of=20 5-Aza-2′-deoxycytidine for 5 days. After this pre-treatment, = the cells were=20 treated with TRAIL or a combination of TRAIL and = 5-Aza-2′-deoxycytidine for=20 additional 72 h. Apoptotic cell death did not change significantly = in TRAIL=20 treated SCGs. Treatment with 5-Aza-2′-deoxycytidine increased the = fraction of=20 apoptotic cells to 16 =B1 5% in NCH421_SCG and to = 13 =B1 5% in=20 NCH441_SCG (Fig. <A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#Fig5">5</A>d).=20 The combination of TRAIL and 5-Aza-2′-deoxycytidine did not = further enhance=20 apoptotic cell death in SCG, indicating that despite the up-regulation = of=20 caspase-8, 5-Aza-2′-deoxycytidine is not sufficient to sensitise = SCGs to=20 TRAIL-induced cell death. </DIV></DIV> <DIV><A name=3DSec21></A> <HR> <DIV class=3Dheading2>Discussion</DIV> In this study we demonstrated that SCGs are resistant to TRAIL = treatment.=20 This finding is in line with previous reports since it has been already = reported=20 that tumour cells, including primary glioma cells, are resistant to the=20 cytotoxic effects of TRAIL [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR19">19</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR20">20</A></CITE>].=20 In addition to TRAIL resistance, glioblastoma cancer stem cells have = been shown=20 to be insensitive to chemotherapy, e.g., temozolomide, carboplatin, = paclitaxel=20 (Taxol) and etoposide (VP16) [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 Also neural stem and progenitor cells were resistant to CD95-induced = cell death,=20 and upon differentiation they became susceptible for CD95-mediated = apoptosis=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR30">30</A></CITE>].=20 In our study we found that the protein synthesis inhibitor = cycloheximide did=20 not sensitise SCGs to TRAIL-mediated cytotoxicity. In contrast, = TRAIL-induced=20 cell death was efficiently enhanced by cycloheximide in NSCGs. Our = results match=20 the data from the literature because it has been reported that = cycloheximide=20 successfully enhanced TRAIL-mediated apoptosis in malignant glioma cell = lines=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR12">12</A></CITE>].=20 Interestingly, cell lines, e.g., U373 glioma cells, that exhibited = suppressed=20 protein levels of caspase-8 could not be sensitised by cycloheximide to=20 TRAIL-mediated apoptosis [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR12">12</A></CITE>],=20 raising the suggestion that caspase-8 might be silenced in SCGs. In line = with=20 this assumption SCGs exhibited low mRNA levels of caspase-8. In = addition, it has=20 been demonstrated that caspase-8 is frequently lost or silenced in human = glioblastoma [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR2">2</A></CITE>]=20 and in a variety of human cancers [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR15">15</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR16">16</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR36">36</A></CITE>].=20 Furthermore, neural stem and progenitor cells known to be resistant to = death=20 receptor-mediated apoptosis induced by CD95 did not express caspase-8 = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR30">30</A></CITE>].=20 However, exogenous or cytokine-mediated expression of caspase-8 was not=20 sufficient to restore their CD95 sensitivity. These authors also found = that in=20 addition to the absence of caspase-8, primitive neural cells expressed = high=20 levels of the death effector domain containing protein PED (also known = as=20 PEA-15), which localised in the DISC and prevented caspase-8 recruitment = and=20 activation [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR30">30</A></CITE>].=20 In the case of neural stem cells, the absence of caspase-8 and high = expression=20 of PED constitute two levels of protection from apoptosis induced by DRs = and=20 inflammatory cytokines in neural stem and progenitor cells. We also = identified a=20 potential mechanism that led to suppression of caspase-8. We found that = the=20 promoter region of CASP8 in SCGs was methylated as demonstrated = by=20 methylation-specific polymerase chain reaction assay. Treatment with=20 5-Aza-2′-deoxycytidine restored caspase-8 protein in both SCG cell = cultures.=20 Conversely, NSCGs had no methylation of the CASP8 promoter. Our results = match=20 published findings in gliomas demonstrating silenced expression of = caspase-8 due=20 to methylation of the CASP8 promoter [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR2">2</A></CITE>].=20 In neuroblastoma, inactivation of CASP8 by hypermethylation is = the=20 hallmark of a defective apoptosis machinery in advanced disease, = suggesting that=20 CASP8 may act as a tumour suppressor gene in this cancer = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR15">15</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR16">16</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR36">36</A></CITE>].=20 In addition, we demonstrated that in SCGs caspase-8 was also = significantly=20 suppressed on the protein level. In previous reports it has been shown = that=20 tumour cells, including primary glioma cells, that are deficient of or = express=20 only low levels of caspase-8 are completely resistant to TRAIL and that=20 induction of caspase-8 is indispensable for TRAIL-induced apoptosis = [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR2">2</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR9">9</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR10">10</A></CITE>].=20 This led us to the conclusion that the suppressed levels of caspase-8 in = SCGs=20 might also explain the high resistance of SCGs to TRAIL-mediated cell = death. We=20 could falsify this hypothesis by restoring caspase-8 expression in both = SCGs by=20 reversing promoter methylation of CASP8 without reversing the = high=20 resistance of these cells towards TRAIL. This contrasts the results of a = previous study in which primary glioma cell lines have been reported to = be=20 resistant to TRAIL stimulation because they expressed low levels of = caspase-8=20 and high levels of the death receptor inhibitor PED/PEA-15 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR7">7</A></CITE>].=20 Treatment with 5-Aza-2′-deoxycytidine resulted in considerable = up-regulation of=20 caspase-8 and sensitisation of primary glioblastoma cells to = TRAIL-induced=20 apoptosis [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR7">7</A></CITE>].=20 Exogenous caspase-8 expression was the main event able to restore TRAIL=20 sensitivity in primary glioblastoma cells. The antitumour activity of=20 5-Aza-2′-deoxycytidine and TRAIL was confirmed in vivo in a mouse = model of=20 glioblastoma multiforme. Evaluation of tumour size, apoptosis and = caspase=20 activation in nude mouse glioblastoma multiforme xenografts showed = dramatic=20 synergy of 5-Aza-2′-deoxycytidine and TRAIL in the treatment of = glioblastoma,=20 whereas the single agents were scarcely effective in terms of reduction = of=20 tumour mass, apoptosis induction and caspase activation. Thus, the = combination=20 of TRAIL and demethylating agents may provide a key tool to overcome=20 glioblastoma resistance to therapeutic treatments. Our results strongly = indicate=20 that in human glioblastoma this combination is insufficient to target = the cancer=20 stem cell fraction. Therefore, TRAIL-based therapies will not fully = eradicate=20 glioblastomas since the cancer stem cell fraction within the tumour is = not=20 efficiently targeted by TRAIL. Hence, treatment strategies are needed = that=20 potently sensitise SCGs to TRAIL-induced cell death. To date, no = treatment=20 strategies specifically sensitising SCGs to TRAIL-mediated cell death = have been=20 described. For primary glioma cell lines such a treatment strategy was = defined=20 and involved the proteasome inhibitor bortezomib [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR21">21</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR22">22</A></CITE>].=20 Bortezomib potently sensitised 13 primary glioma cultures to = TRAIL-mediated=20 apoptosis [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR21">21</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR22">22</A></CITE>].=20 One potential limitation of this study was that they did not demonstrate = that=20 bortezomib and TRAIL also induced apoptosis and synergized in vivo. We also analysed c-FLIP protein, an inhibitor of death = receptor-mediated=20 apoptosis, and detected no difference between SCG and NSCG cells. This = contrasts=20 the results of a recent study in which it was demonstrated that c-FLIP = is=20 significantly up-regulated in CD133+ Jurkat (lymphoma) and CD133+ MCF-7 = (breast=20 carcinoma) cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR40">40</A></CITE>].=20 Down-regulation of FLIP by specific siRNA overcame TRAIL resistance in = CD133+=20 Jurkat and MCF-7 cells, suggesting a major role for c-FLIP in mediating = TRAIL=20 resistance in CD133+ cancer cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR40">40</A></CITE>].=20 However, caspase-8 was not silenced or deleted in CD133+ positive Jurkat = and=20 MCF-7 cells. To identify further factors that possibly convey TRAIL resistance we = employed=20 a proteome array consisting of 30 apoptosis-related proteins. = Interestingly only=20 2 of the 30 proteins were differentially expressed in NSCG and SCG in = both=20 paired cultures, FAS/TNFSF6 and phospho-p53, both of which are not = directly=20 involved in TRAIL-mediated apoptosis. However, in one of the NSCG we = found that=20 TRAIL R2/DR5 was up-regulated. Up-regulation of DR5 has been reported to = confer=20 higher sensitivity to TRAIL [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR1">1</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR29">29</A></CITE>].=20 Recently, it has been demonstrated that the proteasome inhibitor PS-341=20 (bortezomib) up-regulated DR5 expression leading to induction of = apoptosis and=20 enhancement of TRAIL-induced apoptosis despite up-regulation of c-FLIP = and=20 survivin expression in human lung cancer cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR25">25</A></CITE>].=20 Furthermore, it has been reported that the farnesyltransferase inhibitor = R115777=20 up-regulated the expression of DR5 and thereby enhanced TRAIL-mediated = apoptosis=20 in human lung cancer cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR29">29</A></CITE>].=20 Conversely, down-regulation of DR5 by specific siRNA has been shown to = confer=20 TRAIL resistance, suggesting a major role for DR5 in the sensitivity of = cancer=20 cells to TRAIL [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR29">29</A></CITE>].=20 The inhibitor of apoptosis protein (IAP) XIAP, known to inhibit = apoptosis at the=20 level of effector caspase-3/7, was significantly reduced in only one = NSCG. In=20 previous reports XIAP has been linked to confer resistance to TRAIL in = various=20 tumour cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR37">37</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR38">38</A></CITE>].=20 Recently, it has been demonstrated that CD133+ SCGs exhibited higher = XIAP mRNA=20 levels than CD133− glioma cells [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 For the smallest of the IAPs, survivin, we detected no difference = between SCG=20 and NSCG, although it has been reported that the mRNA level of survivin = in=20 CD133+ positive SCGs was significantly higher than in CD133− = glioma cells=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/p536n18xr82w5576/fulltext.htm= l#CR24">24</A></CITE>].=20 Taken together, SCGs were highly resistant towards TRAIL, and this=20 coincidenced with silenced expression of caspase-8, a factor known to be = indispensable for TRAIL-mediated cell death. Reversing methylation of = the=20 caspase-8 promoter led to up-regulation of caspase-8, but failed to = sensitise=20 SCGs to TRAIL-mediated apoptosis. We conclude that whilst treatment with = demethylating agents such as 5-Aza-2′-deoxycytidine sensitises the = bulk tumour=20 cells of glioblastoma, this strategy fails to break the resistance of = CD133+ SCG=20 cells towards TRAIL-mediated apoptosis. As increasing evidence implies = the=20 importance of targeting SCG for an enduring control of glioblastoma, = future=20 research should concentrate on SCG specific sensitisers for = TRAIL-mediated=20 apoptosis. </DIV> <DIV class=3DAcknowledgments><SPAN=20 class=3DAcknowledgmentsHeading>Acknowledgments We= thank=20 Volker Ehemann for performing flow cytometry. 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