10.1007/s11060-009-9967-4</TIT= LE> <META http-equiv=3DContent-Type content=3D"text/html; = charset=3DISO-8859-1"><LINK=20 href=3D"http://www.springerlink.com/config/css/A++_common.css" = type=3Dtext/css=20 rel=3DstyleSheet><LINK=20 href=3D"http://www.springerlink.com/config/css/content_A++_SpringerLink.c= ss"=20 type=3Dtext/css rel=3DstyleSheet> <META content=3D"MSHTML 6.00.6000.16890" name=3DGENERATOR></HEAD> <BODY class=3Dfor-print> <TABLE cellPadding=3D2 border=3D1> <TBODY> <TR class=3Dheader> <TD>Journal of Neuro-Oncology</TD></TR> <TR> <TD>=A9 The Author(s) 2009</TD></TR> <TR> <TD>10.1007/s11060-009-9967-4</TD></TR></TBODY></TABLE> <H2 class=3Drubric>Laboratory Investigation - Human/Animal Tissue</H2> <DIV class=3DHeading1><A name=3Dtitle></A>Different molecular patterns = in=20 glioblastoma multiforme subtypes upon recurrence </DIV> Ramon Martinez1 <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#ContactOfAuthor1"><IMG=20 alt=3D"Contact Information"=20 src=3D"http://www.springerlink.com/content/j12217q235065261/contact.gif" = border=3D0></A>, Veit Rohde1 and=20 Gabriele Schackert2 <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff1></A>(1) </TD> <TD>Department of Neurosurgery, University = of=20 G=F6ttingen, Robert-Koch-Str. 40, 37075 G=F6ttingen,=20 Germany</TD></TR></TBODY></TABLE> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff2></A>(2) </TD> <TD>Department of Neurosurgery, University = of=20 Dresden, Fetscherstr. 74, 01307 Dresden,=20 Germany</TD></TR></TBODY></TABLE> <A name=3DContactOfAuthor1></A> <TABLE class=3DContact> <TBODY> <TR> <TD vAlign=3Dtop><IMG alt=3D"Contact Information"=20 = src=3D"http://www.springerlink.com/content/j12217q235065261/contact.gif" = border=3D0></TD> = <TD>Ramon Martinez</STRONG= > Email:=20 <A=20 = href=3D"mailto:ramon.martinez@med.uni-goettingen.de">ramon.martinez@med.u= ni-goettingen.de</A></TD></TR></TBODY></TABLE> Received:=20 13 April 2009 Accepted:=20 6 July 2009 Published online:=20 31 July 2009 <DIV class=3DAbstract><A name=3DAbs1></A><SPAN=20 class=3DAbstractHeading>Abstract One of the hallmarks = of=20 glioblastoma is its inherent tendency to recur. At this point patients = with=20 relapsed GBM show a survival time of only few months. The molecular = basis of the=20 recurrence process in GBM is still poorly understood. The aim of the = present=20 study was to investigate the genetic profile of relapsed GBM compared to = their=20 respective primary tumors. We have included 20 paired GBMs. In all tumor = samples, we have analyzed p53 and PTEN status by = sequencing=20 analysis, EGFR amplification by semiquantitative PCR and a = wide-genome=20 fingerprinting was performed by microsatellite analysis. Among primary = GBM, we=20 observed twelve type 2 GBM, four type 1 GBM and four further GBM showing = neither=20 p53 mutations nor EGFR amplification (non-type = 1=96non-type 2 GBM).=20 Upon recurrence, we have detected two molecular patterns of tumor = progression:=20 GBM initially showing either type 1 or type 2 profiles conserved them at = the=20 time of relapse. In contrast, non-type 1=96non-type 2 GBM acquired the = typical=20 pattern of type 2 GBM and harbor EGFR amplification without = p53=20 mutation. New PTEN mutations upon relapse were only detected in = type 2=20 GBM. Additional LOH were more frequently identified in relapses of type = 2 GBM=20 than in those showing the type 1 signature. Taken together, our results = strongly=20 suggest that recurrences of GBM may display two distinct pattern of = accumulation=20 of molecular alterations depending on the profile of the original tumor. = </DIV> <DIV class=3DAbstractSection>Electronic=20 supplementary material The online = version of=20 this article (doi:<A=20 href=3D"http://dx.doi.org/10.1007/s11060-009-9967-4">10.1007/s11060-009-9= 967-4</A>)=20 contains supplementary material, which is available to authorized=20 users.</DIV> <SPAN=20 class=3DKeywordHeading>Keywords Glioblastoma -&nbs= p;Relapse - =20 p53 - PTEN=20 - Microsatellite - Mutation <DIV class=3DKeyword><SPAN=20 class=3DKeywordHeading>Abbreviations <SPAN=20 class=3DTerm>GBM Glioblastoma multiforme - <SPAN=20 class=3DTerm>TTR Time to tumor relapse - <SPAN=20 class=3DTerm>MGMT O 6=20 -methyl-guanine-methyl-transferase - <SPAN=20 class=3DTerm>CFTR Cystic fibrosis transmembrane regulator - = <SPAN=20 class=3DTerm>KPS Karnofsky performance score </DIV> <DIV class=3DFulltext> <DIV class=3D""><A name=3DSec1></A> <HR> <DIV class=3Dheading2>Introduction</DIV> Glioblastoma multiforme is the most frequent primary brain = tumor in=20 adults. One of its characteristic features is the intrinsic tendency to = recur=20 despite aggressive therapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR1">1</A></CITE>].=20 Upon recurrence, the median survival of patients is only few months = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR2">2</A></CITE>]=20 whereas the overall median survival time is 15 months. This has not = significantly changed in the last 20 years despite of advances in = surgery,=20 radio- und chemotherapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR4">4</A></CITE>]=20 A moderate improvement of the 2-year survival rate has been achieved = after=20 therapy with the alkylating drug temozolomide in GBM with = hypermethylated=20 MGMT compared to those with unmethylated gene (46% vs. 26%) = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR5">5</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR6">6</A></CITE>].=20 GBM is one of the most intensively investigated human = malignancy but=20 the molecular mechanisms associated with recurrence are still poorly = understood.=20 Saxena et al. [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR7">7</A></CITE>]=20 reported a high rate of homozygous deletion of the genes p16=20 INK4 and p15 INK4a in GBM = relapses.=20 In a further report, this author investigated 10 GBM and their = corresponding=20 relapses and observed that recurrences displayed the pattern of genetic=20 alterations frequently observed in the de novo pathway (GBM without a = low grade=20 precursor lesion) [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR8">8</A></CITE>].=20 Hulsebos et al. [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR9">9</A></CITE>]=20 analyzed 12 match paired GBM and found in relapse cases new LOH at = chromosome=20 regions 1p36, 19q13, 10q23 and 1q25. In contrast to the former study, = GBM=20 relapses encompassing new p53 mutation or EGFR = amplification were=20 not observed. The identification of molecular features associated with = recurrences=20 of GBM is of major importance, since a better understanding of this = process=20 might provide clues for the development of efficient treatments. In the = present=20 analysis, we have examined the molecular signatures of forty tumors = consisting=20 in relapsed GBM and their corresponding primary neoplasms which had = undergone=20 surgery and radio-chemotherapy. This included the status of p53,=20 PTEN and EGFR as well as a wide genome LOH analysis with = thirteen=20 highly informative microsatellite markers at chromosome regions 17p13, = 10q23,=20 1p35-36, 19q13, 13q14, 9p21. </DIV> <DIV class=3D""><A name=3DSec2></A> <HR> <DIV class=3Dheading2>Materials and methods</DIV> <DIV class=3D""><A name=3DSec3></A> <DIV class=3DHeading3>Patient population and tumor samples</DIV> All patients had undergone surgery with the goal of = maximal possible=20 tumor resection followed by fractionated radiotherapy (mean dose: 58 = Gray) and=20 chemotherapy. Forty paired tumors from twenty patients were available. = Informed=20 consent for samples and data analysis from each patient or the = patient=92s=20 caretaker was obtained. Tumor samples were immediately frozen in liquid nitrogen = after=20 surgical resection and stored at −80=B0C. Tumor tissue was = evaluated by=20 experienced pathologists according to the 2000 WHO classification = criteria. DNA=20 from tumor specimens was isolated applying the QIAamp=AE DNA = Mini Kit=20 (Qiagen, Hilden, Germany). </DIV> <DIV class=3D""><A name=3DSec4></A> <DIV class=3DHeading3>LOH analysis</DIV> LOH of chromosome 10q was studied with markers covering = reported=20 deletions on 10q23-24: D10S215, D10S541 and the intragenic PTEN = marker=20 PTENCA. Analysis of LOH at 17p13 was performed with a p53 = intragenic=20 localized marker. Allelic losses at 9p21 (p16 INK4a = ,=20 p14 ARF and p15) and 13q14 (RB1) = were=20 studied with primers D9S171, D9S1748, D9S1749, D13S153 and D13S267. Loss = of=20 heterozygosity at 1p35-p36 and 19q13 was assessed with the markers = D1S468,=20 D1S482, D19S112 and D19S412. Amplified PCR products were analyzed on a=20 denaturing 6.5% Long Ranger polyacrylamide gel on an Automated Laser=20 Fluorescence sequencing device and analyzed using the = ALLELELINKS=AE=20 1.00 software (Amersham Pharmacia Biotech, Freiburg, Germany). = Evaluation of LOH=20 was performed as described [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR10">10</A></CITE>].=20 </DIV> <DIV class=3D""><A name=3DSec5></A> <DIV class=3DHeading3>Semiquantitative PCR-analysis</DIV> For studying amplification of EGFR a differential = duplex-PCR=20 with the CFTR gene marker was carried out, as described [<CITE><A = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR11">11</A></CITE>].=20 Briefly, we calculated the EGFR/CFTR ratios [x] from = peripheral=20 blood DNA of 20 healthy Caucasian adults. A value = [x + 3SD] was=20 considered as the cut-off level for the normal gene copy number. Ratios = higher=20 than [x + 3SD] were regarded as evidence of more than 2 = copies=20 of the EGFR gene. </DIV> <DIV class=3D""><A name=3DSec6></A> <DIV class=3DHeading3>Sequencing analysis of p53 and PTEN</DIV> Mutational analysis of p53 and PTEN were = performed in=20 the DNA from both tumors and corresponding leukocytes. After amplifying = all=20 exons and intron-exon boundaries, we analyzed the PCR products on 1% = agarose gel=20 and excised and eluted them in sterile H2O. Subsequently, = they were=20 subjected to cycle sequencing reactions using the Thermo = Sequenase=AE=20 Fluorescent Cycle Sequencing kit (Amersham Pharmacia Biotech). The cycle = sequencing products were resolved using a denaturing 6.5% Long Ranger=20 polyacrylamide gel on a sequencing device. PTEN primer sequences = and PCR=20 conditions were previously described [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR12">12</A></CITE>].=20 P53 primer sequences and PCR conditions are based on genome = database=20 entries (<A=20 href=3D"http://www.ncbi.nlm.nih.gov/">http://www.ncbi.nlm.nih.gov/</A>). = Briefly,=20 PCR reactions were carried out using a mixture containing 150 ng = DNA,=20 10 mM Tris, 2 mM MgCl2, HCl (pH 8.3), 0.2 mM = dNTP,=20 10 mM concentrations of each primer and 0.5 unit of Taq DNA = Polymerase.=20 Temperature cycles and times for PCR reactions were: denaturation at = 94=B0C for=20 30 s, annealing at 65=B0C for 30 s, and extension at 72=B0C = for 30 s.=20 Each PCR reaction was preceded by a 3-min denaturation at 94=B0C, and = the final=20 cycle was followed by a 3-min extension at 72=B0C. The total number of = cycles for=20 PCR amplification was 25=9630, depending on the sample DNA. </DIV> <DIV class=3D""><A name=3DSec7></A> <DIV class=3DHeading3>Statistical analysis</DIV> Student t-test, Chi-square test, Fisher=92s exact = test and=20 Mann=96Whitney U-test, were performed to compare differences between = groups=20 depending on the analyzed variables. Spearman correlation test was used = to=20 evaluate the correlation between two parameters. Confidence interval = (CI) was=20 obtained through logistic regression. Survival studies were performed = with the=20 Kaplan Meier analysis and the log-rank test. A value of=20 P < 0.05 was considered to be significant. Analyses = were=20 performed with the SPSS software (version 10, SPSS Inc., Chicago, IL, = USA).=20 </DIV></DIV> <DIV class=3D""><A name=3DSec8></A> <HR> <DIV class=3Dheading2>Results</DIV> <DIV class=3D""><A name=3DSec9></A> <DIV class=3DHeading3>Clinico-pathological characteristics of the = patients</DIV> The male/female ratio was 1:1. The median age at diagnosis = of=20 primary tumor was 59.5 years (range: 27=9669; SD: 12.25). The = median time to=20 tumor recurrence was 7.5 months (range: 3=9624 months; SD: = 5.1; 95%-CI:=20 5.3=967.4 months). With one only exception, all patients displayed = at=20 diagnosis a Karnofsky performance score (KPS) of >70 points. An = association=20 between age or KPS and TTR was not evidenced = (P =3D 0.35,=20 ρ =3D 0.30, Spearman correlation test). There were no = significant=20 associations between mutation of p53 or EGFR amplification = and=20 either survival time (P =3D 0.250 and 0.127, = respectively,=20 log-rank test and Kaplan=96Meier) or time to tumor relapse=20 (P =3D 0.210 and 0.287, respectively, log-rank test and = Kaplan=96Meier). </DIV> <DIV class=3D""><A name=3DSec10></A> <DIV class=3DHeading3>Genetic analyses</DIV> <DIV class=3DPara> <DIV class=3D"">In primary GBM we have observed EGFR = amplification without=20 mutations of p53 in 12 cases, thus indicating a type II GBM = profile. Four=20 further cases showed p53 mutations (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab1">1</A>,=20 Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig1">1</A>)=20 in the absence of EGFR amplification, which was indicative of a = type 1=20 GBM signature. In four GBM neither p53 mutations nor EGFR=20 amplification were identified (non-type 1=96non-type 2 GBM).<A = name=3DTab1></A> <DIV class=3DCapt>Table 1 Sequence = variations=20 of p53 in type 1 glioblastomas </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH align=3Dleft colSpan=3D6> p53 sequence variations </TH></TR> <TR class=3Dheader> <TH align=3Dleft> Case</TH> <TH align=3Dleft> Sequence variation</TH> <TH align=3Dleft> Base exchange</TH> <TH align=3Dleft> Aminoacid substitution</TH> <TH align=3Dleft> CpG site</TH> <TH align=3Dleft> Distribution</TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> G3p</TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.266G>T</TD> <TD align=3Dleft> GGA→GTA</TD> <TD align=3Dleft> Gly→Val</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 44 tumors No germline mutationsa </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G3r</TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.266G>T</TD> <TD align=3Dleft> GGA→GTA</TD> <TD align=3Dleft> Gly→Val</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 44 tumors No germline mutationsa </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G7p</TD> <TD align=3Dleft> c.72C>G (Arg72Pro)</TD> <TD align=3Dleft> CCC→CGC</TD> <TD align=3Dleft> Arg→Pro</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Validated polymorphism Higher risk lung cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR34">34</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS3+21_37dup (P53PIN3) </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Higher risk of CRC [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR36">36</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G7r</TD> <TD align=3Dleft> c.72C>G (Arg72Pro)</TD> <TD align=3Dleft> CCC→CGC</TD> <TD align=3Dleft> Arg→Pro</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Validated polymorphism Higher risk lung cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR34">34</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS3+21_37dup (P53PIN3) </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphismaHigher risk of CRC = [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR36">36</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G14p</TD> <TD align=3Dleft> c.14G>C</TD> <TD align=3Dleft> CTG→CTC</TD> <TD align=3Dleft> Leu→Leu</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G14r</TD> <TD align=3Dleft> c.14G>C</TD> <TD align=3Dleft> CTG→CTC</TD> <TD align=3Dleft> Leu→Leu</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G16p</TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS2+38G>C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR41">41</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G16r</TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS2+38G>C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR41">41</A></CITE>]=20 </TD></TR></TBODY></TABLE> <DIV class=3DCapt> <DIV class=3DCaptCont> <DIV class=3D"">aAccording to p53 data bases entries, = mut=20 mutation, CRC colorectal cancer </DIV></DIV></DIV> <DIV class=3DFigure><A name=3DFig1></A><IMG=20 alt=3DMediaObjects/11060_2009_9967_Fig1_HTML.gif=20 src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Fig1_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 1 Upper = row:=20 wild type sequence of p53. Lower row: base exchange = mutation=20 (GGA→GTA) of p53 (exon 8, codon 266), predicting the = aminoacid=20 substitution Gly→Val. The arrow indicates the base exchange </DIV> <HR> </DIV></DIV> <DIV class=3DPara> <DIV class=3D"">At the time of recurrence, every type 1 GBM and type 2 = GBM=20 conserved their genetic pattern concerning p53 mutational status = and=20 EGFR amplification. On the other hand, relapses of non-type = 1=96non-type 2=20 GBM showed in all cases a new EGFR amplification, thus acquiring = the=20 signature of type 2 GBM. Additional PTEN mutations in relapse = cases were=20 observed only among type 2 GBM (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab2">2</A>).=20 Overall, additional LOH were observed more frequently in relapses of = type 2 GBM=20 (83.3%), than in those of type 1 (50%) or non-type 1=96non-type 2 tumors = (25%).=20 Further results are shown in Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>.<A=20 name=3DTab2></A> <DIV class=3DCapt>Table 2 Sequence = variations=20 of PTEN in first and relapse glioblastomas </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH align=3Dleft colSpan=3D5> PTEN sequence variations </TH></TR> <TR class=3Dheader> <TH align=3Dleft> Case</TH> <TH align=3Dleft> Sequence variation</TH> <TH align=3Dleft> Mutation effect</TH> <TH align=3Dleft> Aminoacid substitution</TH> <TH align=3Dleft> Distribution</TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> G1p</TD> <TD align=3Dleft> c.44C>T</TD> <TD align=3Dleft> GGC→GGT</TD> <TD align=3Dleft> Gly→Gly</TD> <TD align=3Dleft> Bladder cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR39">39</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G1r</TD> <TD align=3Dleft> c.44C>T</TD> <TD align=3Dleft> GGC→GGT</TD> <TD align=3Dleft> Gly→Gly</TD> <TD align=3Dleft> Bladder cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR39">39</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G2p</TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G2r</TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G4</TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G4r</TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G5p</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS2-96A>G</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G5r</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.88A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Head neck squamous cell cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR42">42</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G11p</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G11r</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.88A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Head & neck squamous cell cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR42">42</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8-2A>G</TD> <TD align=3Dleft> Splice mutation</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G13p</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G13r</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8-2A>G</TD> <TD align=3Dleft> Splice mutation</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.75C>A</TD> <TD align=3Dleft> CAT→AAT</TD> <TD align=3Dleft> His→Asn</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G17p</TD> <TD align=3Dleft> IVS4 109_110 ins TCTTA</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G17r</TD> <TD align=3Dleft> IVS4 109_110 ins TCTTA</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G19p</TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G19r</TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.252A>G</TD> <TD align=3Dleft> GAT→GGT</TD> <TD align=3Dleft> Asp→Gly</TD> <TD align=3Dleft> Glioma [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR43">43</A></CITE>]=20 </TD></TR></TBODY></TABLE><A name=3DTab3></A> <DIV class=3DCapt>Table 3 Review = of genetic=20 changes of primary and relapsed glioblastomas </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft colSpan=3D2> <IMG = alt=3DMediaObjects/11060_2009_9967_Taba_HTML.gif=20 = src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Taba_HTML.gif"=20 align=3Dmiddle> </TD></TR></TBODY></TABLE> <DIV class=3DCapt> <DIV class=3DCaptCont> <DIV class=3D"">TTR time to tumor relapse (months), Mut = mutation,=20 Ampl amplification, p primary (first tumor), r = relapse=20 (first relapse), m male, f female, no ther no = adjuvant=20 therapy performed, RT radiotherapy, A ACNU (nimustine), = V=20 VM26 (teniposide), T temozolomide. Black boxes loss of=20 heterozygosity, grey boxes retention of heterozygosity, white=20 boxes not informative marker </DIV></DIV> <DIV class=3DCaptCont> <DIV class=3D"">* Age (years) at first=20 operation</DIV></DIV></DIV></DIV></DIV> Among p53 mutations there were one C:G to A:T = transition=20 mutation at CpG site (codon 273), one C>T transition at non-CpG site = (codon=20 278), two transversion mutations at non-CpG sites (G>C at codon 14 = and G>T=20 at codon 266, Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig1">1</A>)=20 as well as one validated polymorphism in exon 4 (G>C at codon 72). = Moreover,=20 we have observed two validated intronic polymorphisms in intron 3=20 (IVS3+21_37dup) and intron 2 (IVS2+38G>C). One of the mutations = (c.14G>C)=20 has not been previously described in p53 data bases (<A=20 href=3D"http://http//www.p53.iarc.fr">http://http//www.p53.iarc.fr</A>, = <A=20 href=3D"http://http//www.p53.free.fr">http://http//www.p53.free.fr</A>). = A=20 detailed characterization of p53 sequence variations is shown in=20 Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab1">1</A>.=20 <DIV class=3DPara> <DIV class=3D"">Concerning PTEN mutations, we have identified = four=20 transition mutations in exons 2, 5, 6 and 7, in all cases predicting = aminoacid=20 substitution. Furthermore, there was an acceptor site splice mutation at = the=20 conserved junction nucleotides (IVS8-2A>G). Finally, we have observed = two=20 transversion mutations in exon 5 (c.105G>T) and exon 4 (c.75C>A,=20 Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig2">2</A>)=20 as well as four intronic polymorphisms (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab2">2</A>).=20 The transition in exon 6 (c.177A>G), the splicing mutation and two of = the=20 polymorphisms have not been previously reported (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab2">2</A>).=20 <DIV class=3DFigure><A name=3DFig2></A><IMG=20 alt=3DMediaObjects/11060_2009_9967_Fig2_HTML.gif=20 src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Fig2_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 2 Upper = row:=20 wild type sequence of PTEN. Lower row: base exchange = mutation=20 (CAT→AAT) of PTEN (exon 4, codon 75) predicting the = aminoacid=20 substitution His→Asn. The arrow indicates the base exchange </DIV> <HR> </DIV></DIV> The results of EGFR amplification analysis and = wide-genome=20 LOH are shown in Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>=20 and Supplementary Table 1 (EGFR/CFTR ratios). Interestingly, = GBM=20 cases harboring mutations of p53 always displayed LOH at 17p13 as = well.=20 Similarly, tumors featuring PTEN mutations showed LOH at 10q23, = at least=20 in one of the microsatellites used to analyze this region. = </DIV></DIV> <DIV class=3D""><A name=3DSec11></A> <HR> <DIV class=3Dheading2>Discussion</DIV> GBM mostly exceed in its occurrence and mortality beyond = any other=20 primary brain tumor in adults [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR13">13</A></CITE>].=20 On molecular level, two subsets of GBM have been recognized on the basis = of=20 genetic make-up and clinical features [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR14">14</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR15">15</A></CITE>].=20 Type 2 (de novo) GBM occurs commonly in elderly patients and exhibit=20 overexpression and/or amplification of EGFR. Type 1 (secondary) = GBM shows=20 typically histological progression from a previous low grade astrocytoma = and=20 affects young patients. Paradigmatically, mutations of p53 will = be=20 observed (for review see [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR15">15</A></CITE>]).=20 Most of GBM cases belong to the type 2 group and genetic events are = mutually=20 exclusive for EGFR and p53. Moreover, there are GBM = without either=20 EGFR amplification or p53 mutations (non-type 1=96non-type = 2 GBM)=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR16">16</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR17">17</A></CITE>].=20 During the past decade, accumulated evidence pointed to that type 1 and = type 2=20 GBM constitute distinct disease entities developing through different = genetic=20 pathways [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR13">13</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR14">14</A></CITE>],=20 showing different RNA and protein patterns [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR18">18</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR19">19</A></CITE>]=20 and also probably differing in response to treatment [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR15">15</A></CITE>].=20 After multimodal therapy, GBM almost always recur. = Although this is=20 a capital feature of GBM, only few investigations with series of match = paired=20 patients are available from the literature and available data is = partially=20 contradictory [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR7">7</A></CITE>=96<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR9">9</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR20">20</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR21">21</A></CITE>].=20 We aimed to investigate whether molecular subtypes of GBM also show = different=20 profiles at the time of relapse. <DIV class=3DPara> <DIV class=3D"">In our genetic analysis, we have observed two patterns = of tumor=20 recurrence (Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig3">3</A>):=20 GBM type 1 and type 2 retained upon recurrence their genetic alterations = affecting p53 and EGFR. In contrast, all relapses of = non-type=20 1=96non-type 2 GBM showed a new EGFR amplification, thus they = acquired a=20 type 2 GBM profile. Accumulation of additional genetic alterations upon = relapse=20 included PTEN mutations in four type 2 GBM (but not in the other=20 subtypes). Additional LOH were more frequently observed in relapses of = type 2=20 GBM (in 10/12 cases) compared to relapses of type 1 GBM and non-type = 1=96non-type=20 2 GBM (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>).=20 The accumulation of alterations during the clonal expansion of = transformed glial=20 cells is well known in GBM, confer tumor cells a growth advantage and = underlies=20 the genetic heterogeneity of GBM [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR22">22</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR23">23</A></CITE>].=20 This phenomenon contributes to explain the molecular differences between = primary=20 tumors and relapses [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR9">9</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR24">24</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR25">25</A></CITE>].=20 <DIV class=3DFigure><A name=3DFig3></A><IMG=20 alt=3DMediaObjects/11060_2009_9967_Fig3_HTML.gif=20 src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Fig3_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 3 Graphic = showing two=20 molecular patterns of GBM recurrence </DIV> <HR> </DIV></DIV> As a part of one previous investigation we had = characterized the=20 promoter methylation status of the DNA repair gene MGMT in all = type 1 GBM=20 (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>,=20 cases G3, G7, G14 and G16), non-type 1=96non-type 2 GBM (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>,=20 cases G2, G6, G8 and G9) as well as eight type 2 GBM here described = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR26">26</A></CITE>].=20 This analysis together with the present results of p53 and = EGFR=20 have evidenced similar rates of MGMT methylation for the three = GBM=20 subtypes, and they are in accordance with the overall methylation rates = reported=20 in glioblastoma [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR5">5</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR27">27</A></CITE>].=20 Typically, MGMT was methylated in the first GBM and remained = always=20 methylated upon relapse. Thus, our results also confirmed the = observation that,=20 in the pathogenesis and progression of GBM, MGMT becomes = hypermethylated=20 at an early stage [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR28">28</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR29">29</A></CITE>]=20 and remains methylated through progression, a situation which was has = been=20 described in small adenomas and carcinomas of colorectal origin as well=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR30">30</A></CITE>].=20 The p53 mutational analysis in those type 1 GBM with methylated=20 MGMT showed G:C to A:T transition within CpG site (c.273 = CGT→CAT) in two=20 cases. Further mutations of p53 in tumors with methylated = MGMT (1=20 case) occurred in non-CpG sites. From p53 data bases entries,=20 approximately 52% of mutational events are transition mutations and, of = this=20 subset, about 72% are G:C to A:T transitions (<A=20 href=3D"http://http//www.p53.iarc.fr">http://http//www.p53.iarc.fr</A>, = <A=20 href=3D"http://http//www.p53.free.fr">http://http//www.p53.free.fr</A>) = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR31">31</A></CITE>],=20 although the signature of the mutational spectrum varies according to = tumor=20 type. It was previously suggested a link [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR29">29</A></CITE>]=20 between MGMT methylation and the presence of G:C to A:T = transition=20 mutations in p53 in gliomas. In the present study we could = confirm the=20 former in two patients (cases G14p, G14r, G16p and G16r, Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab1">1</A>);=20 in these cases, G:C to A:T transitions could be attributable to a = inactivation=20 in MGMT that allows O 6 -methylguanine = to=20 persist and be read as an adenine. Our observations contrast with a = recent=20 investigation about GBM reporting only 4% of mutations of p53 in = tumors=20 with methylated MGMT within CpG sites [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR32">32</A></CITE>]=20 although we should take in mind the size of our type 1 GBM collective, = clearly=20 too short to state a definite tendency. Strikingly, we have identified in cases G7p and G7r the = exonic=20 p53 polymorphism c.72C>G (Arg72Pro) together with the intronic = polymorphism IVS3+21_37dup (P53PIN3). Several investigations have = pointed=20 the association between p53 polymorphisms and an increased risk = for=20 different cancers: for instance, in the case of the exonic c.72 variant = such=20 association was suggested since the p53 protein with Arg72 is more = efficient in=20 inducing apoptosis than the one with the Pro72 variant [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR33">33</A></CITE>].=20 In clinical studies, the variants c.72 Pro/Pro and c.72 Arg/Pro have = been=20 associated with an elevated risk of lung cancer [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR34">34</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR35">35</A></CITE>].=20 Recently, intron 3 duplication (IVS3+21_37dup) has been found to be = associated=20 with increased risk of colorectal cancer [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR36">36</A></CITE>].=20 However, the authors could not determine whether this intron 3 = duplication alone=20 influences mRNA stability or if this effect requires the codon Pro72 = variant. To=20 our knowledge, the occurrence of both polymorphisms in GBM patients has = not been=20 reported before and pointed to a possible role in brain cancer as well. = With respect to PTEN, we have identified mutations = only in=20 type 2 GBM. In four cases, PTEN mutations occurred in both the = first GBM=20 and the relapse. In four further cases, new mutations were observed in = the=20 recurrences but not in the first tumor. The rate and pattern of = mutations are in=20 accordance with previous reported data [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR37">37</A></CITE>].=20 All tumors with PTEN mutations also harbored LOH at the = chromosome region=20 10q23, which pointed to the inactivation of this tumor suppressor gene = and make=20 patent its contribution to the tumorigenesis and progression in GBM. = Taken together, our results strongly suggest that GBM = relapses,=20 compared to the corresponding first tumor, accumulate additional = molecular=20 alterations and may develop along two distinct pathways, which = contribute to=20 delineate the different profiles of type 1, type 2 and non-type = 1=96non-type 2=20 GBM. </DIV> <DIV class=3DAcknowledgments>Open=20 Access This article is distributed = under the=20 terms of the Creative Commons Attribution Noncommercial License which = permits=20 any noncommercial use, distribution, and reproduction in any medium, = provided=20 the original author(s) and source are credited.</DIV> <DIV class=3DAppendix><A name=3DAppESM1></A> <DIV class=3D""><A name=3DSecESM1></A> <HR> <DIV class=3Dheading2>Electronic supplementary material</DIV> <DIV class=3DPara> <DIV class=3D"">Below is the link to the electronic supplementary = material.=20 <DIV><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects= /11060_2009_9967_MOESM1_ESM.pdf"> <DIV class=3DCapt>Supplementary material 1 (PDF 25=20 kb)</DIV></A></DIV></DIV></DIV></DIV></DIV> <HR> <H2><A name=3DBib1></A>References </H2> <TABLE> <TBODY class=3DCitation> <TR vAlign=3Dtop> <TD>1.</TD> <TD><A name=3DCR1></A>Salcman M, Kaplan R (1991) Intracranial tumors = in=20 adults. 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From: Subject: 10.1007/s11060-009-9967-4 Date: Sat, 29 Aug 2009 14:09:09 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_0002_01CA28B2.470D8AC0" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.5579 This is a multi-part message in MIME format. ------=_NextPart_000_0002_01CA28B2.470D8AC0 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.springerlink.com/content/j12217q235065261/fulltext.html 10.1007/s11060-009-9967-4</TIT= LE> <META http-equiv=3DContent-Type content=3D"text/html; = charset=3DISO-8859-1"><LINK=20 href=3D"http://www.springerlink.com/config/css/A++_common.css" = type=3Dtext/css=20 rel=3DstyleSheet><LINK=20 href=3D"http://www.springerlink.com/config/css/content_A++_SpringerLink.c= ss"=20 type=3Dtext/css rel=3DstyleSheet> <META content=3D"MSHTML 6.00.6000.16890" name=3DGENERATOR></HEAD> <BODY class=3Dfor-print> <TABLE cellPadding=3D2 border=3D1> <TBODY> <TR class=3Dheader> <TD>Journal of Neuro-Oncology</TD></TR> <TR> <TD>=A9 The Author(s) 2009</TD></TR> <TR> <TD>10.1007/s11060-009-9967-4</TD></TR></TBODY></TABLE> <H2 class=3Drubric>Laboratory Investigation - Human/Animal Tissue</H2> <DIV class=3DHeading1><A name=3Dtitle></A>Different molecular patterns = in=20 glioblastoma multiforme subtypes upon recurrence </DIV> Ramon Martinez1 <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#ContactOfAuthor1"><IMG=20 alt=3D"Contact Information"=20 src=3D"http://www.springerlink.com/content/j12217q235065261/contact.gif" = border=3D0></A>, Veit Rohde1 and=20 Gabriele Schackert2 <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff1></A>(1) </TD> <TD>Department of Neurosurgery, University = of=20 G=F6ttingen, Robert-Koch-Str. 40, 37075 G=F6ttingen,=20 Germany</TD></TR></TBODY></TABLE> <TABLE> <TBODY> <TR vAlign=3Dtop> <TD><A = name=3DAff2></A>(2) </TD> <TD>Department of Neurosurgery, University = of=20 Dresden, Fetscherstr. 74, 01307 Dresden,=20 Germany</TD></TR></TBODY></TABLE> <A name=3DContactOfAuthor1></A> <TABLE class=3DContact> <TBODY> <TR> <TD vAlign=3Dtop><IMG alt=3D"Contact Information"=20 = src=3D"http://www.springerlink.com/content/j12217q235065261/contact.gif" = border=3D0></TD> = <TD>Ramon Martinez</STRONG= > Email:=20 <A=20 = href=3D"mailto:ramon.martinez@med.uni-goettingen.de">ramon.martinez@med.u= ni-goettingen.de</A></TD></TR></TBODY></TABLE> Received:=20 13 April 2009 Accepted:=20 6 July 2009 Published online:=20 31 July 2009 <DIV class=3DAbstract><A name=3DAbs1></A><SPAN=20 class=3DAbstractHeading>Abstract One of the hallmarks = of=20 glioblastoma is its inherent tendency to recur. At this point patients = with=20 relapsed GBM show a survival time of only few months. The molecular = basis of the=20 recurrence process in GBM is still poorly understood. The aim of the = present=20 study was to investigate the genetic profile of relapsed GBM compared to = their=20 respective primary tumors. We have included 20 paired GBMs. In all tumor = samples, we have analyzed p53 and PTEN status by = sequencing=20 analysis, EGFR amplification by semiquantitative PCR and a = wide-genome=20 fingerprinting was performed by microsatellite analysis. Among primary = GBM, we=20 observed twelve type 2 GBM, four type 1 GBM and four further GBM showing = neither=20 p53 mutations nor EGFR amplification (non-type = 1=96non-type 2 GBM).=20 Upon recurrence, we have detected two molecular patterns of tumor = progression:=20 GBM initially showing either type 1 or type 2 profiles conserved them at = the=20 time of relapse. In contrast, non-type 1=96non-type 2 GBM acquired the = typical=20 pattern of type 2 GBM and harbor EGFR amplification without = p53=20 mutation. New PTEN mutations upon relapse were only detected in = type 2=20 GBM. Additional LOH were more frequently identified in relapses of type = 2 GBM=20 than in those showing the type 1 signature. Taken together, our results = strongly=20 suggest that recurrences of GBM may display two distinct pattern of = accumulation=20 of molecular alterations depending on the profile of the original tumor. = </DIV> <DIV class=3DAbstractSection>Electronic=20 supplementary material The online = version of=20 this article (doi:<A=20 href=3D"http://dx.doi.org/10.1007/s11060-009-9967-4">10.1007/s11060-009-9= 967-4</A>)=20 contains supplementary material, which is available to authorized=20 users.</DIV> <SPAN=20 class=3DKeywordHeading>Keywords Glioblastoma -&nbs= p;Relapse - =20 p53 - PTEN=20 - Microsatellite - Mutation <DIV class=3DKeyword><SPAN=20 class=3DKeywordHeading>Abbreviations <SPAN=20 class=3DTerm>GBM Glioblastoma multiforme - <SPAN=20 class=3DTerm>TTR Time to tumor relapse - <SPAN=20 class=3DTerm>MGMT O 6=20 -methyl-guanine-methyl-transferase - <SPAN=20 class=3DTerm>CFTR Cystic fibrosis transmembrane regulator - = <SPAN=20 class=3DTerm>KPS Karnofsky performance score </DIV> <DIV class=3DFulltext> <DIV class=3D""><A name=3DSec1></A> <HR> <DIV class=3Dheading2>Introduction</DIV> Glioblastoma multiforme is the most frequent primary brain = tumor in=20 adults. One of its characteristic features is the intrinsic tendency to = recur=20 despite aggressive therapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR1">1</A></CITE>].=20 Upon recurrence, the median survival of patients is only few months = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR2">2</A></CITE>]=20 whereas the overall median survival time is 15 months. This has not = significantly changed in the last 20 years despite of advances in = surgery,=20 radio- und chemotherapy [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR4">4</A></CITE>]=20 A moderate improvement of the 2-year survival rate has been achieved = after=20 therapy with the alkylating drug temozolomide in GBM with = hypermethylated=20 MGMT compared to those with unmethylated gene (46% vs. 26%) = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR3">3</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR5">5</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR6">6</A></CITE>].=20 GBM is one of the most intensively investigated human = malignancy but=20 the molecular mechanisms associated with recurrence are still poorly = understood.=20 Saxena et al. [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR7">7</A></CITE>]=20 reported a high rate of homozygous deletion of the genes p16=20 INK4 and p15 INK4a in GBM = relapses.=20 In a further report, this author investigated 10 GBM and their = corresponding=20 relapses and observed that recurrences displayed the pattern of genetic=20 alterations frequently observed in the de novo pathway (GBM without a = low grade=20 precursor lesion) [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR8">8</A></CITE>].=20 Hulsebos et al. [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR9">9</A></CITE>]=20 analyzed 12 match paired GBM and found in relapse cases new LOH at = chromosome=20 regions 1p36, 19q13, 10q23 and 1q25. In contrast to the former study, = GBM=20 relapses encompassing new p53 mutation or EGFR = amplification were=20 not observed. The identification of molecular features associated with = recurrences=20 of GBM is of major importance, since a better understanding of this = process=20 might provide clues for the development of efficient treatments. In the = present=20 analysis, we have examined the molecular signatures of forty tumors = consisting=20 in relapsed GBM and their corresponding primary neoplasms which had = undergone=20 surgery and radio-chemotherapy. This included the status of p53,=20 PTEN and EGFR as well as a wide genome LOH analysis with = thirteen=20 highly informative microsatellite markers at chromosome regions 17p13, = 10q23,=20 1p35-36, 19q13, 13q14, 9p21. </DIV> <DIV class=3D""><A name=3DSec2></A> <HR> <DIV class=3Dheading2>Materials and methods</DIV> <DIV class=3D""><A name=3DSec3></A> <DIV class=3DHeading3>Patient population and tumor samples</DIV> All patients had undergone surgery with the goal of = maximal possible=20 tumor resection followed by fractionated radiotherapy (mean dose: 58 = Gray) and=20 chemotherapy. Forty paired tumors from twenty patients were available. = Informed=20 consent for samples and data analysis from each patient or the = patient=92s=20 caretaker was obtained. Tumor samples were immediately frozen in liquid nitrogen = after=20 surgical resection and stored at −80=B0C. Tumor tissue was = evaluated by=20 experienced pathologists according to the 2000 WHO classification = criteria. DNA=20 from tumor specimens was isolated applying the QIAamp=AE DNA = Mini Kit=20 (Qiagen, Hilden, Germany). </DIV> <DIV class=3D""><A name=3DSec4></A> <DIV class=3DHeading3>LOH analysis</DIV> LOH of chromosome 10q was studied with markers covering = reported=20 deletions on 10q23-24: D10S215, D10S541 and the intragenic PTEN = marker=20 PTENCA. Analysis of LOH at 17p13 was performed with a p53 = intragenic=20 localized marker. Allelic losses at 9p21 (p16 INK4a = ,=20 p14 ARF and p15) and 13q14 (RB1) = were=20 studied with primers D9S171, D9S1748, D9S1749, D13S153 and D13S267. Loss = of=20 heterozygosity at 1p35-p36 and 19q13 was assessed with the markers = D1S468,=20 D1S482, D19S112 and D19S412. Amplified PCR products were analyzed on a=20 denaturing 6.5% Long Ranger polyacrylamide gel on an Automated Laser=20 Fluorescence sequencing device and analyzed using the = ALLELELINKS=AE=20 1.00 software (Amersham Pharmacia Biotech, Freiburg, Germany). = Evaluation of LOH=20 was performed as described [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR10">10</A></CITE>].=20 </DIV> <DIV class=3D""><A name=3DSec5></A> <DIV class=3DHeading3>Semiquantitative PCR-analysis</DIV> For studying amplification of EGFR a differential = duplex-PCR=20 with the CFTR gene marker was carried out, as described [<CITE><A = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR11">11</A></CITE>].=20 Briefly, we calculated the EGFR/CFTR ratios [x] from = peripheral=20 blood DNA of 20 healthy Caucasian adults. A value = [x + 3SD] was=20 considered as the cut-off level for the normal gene copy number. Ratios = higher=20 than [x + 3SD] were regarded as evidence of more than 2 = copies=20 of the EGFR gene. </DIV> <DIV class=3D""><A name=3DSec6></A> <DIV class=3DHeading3>Sequencing analysis of p53 and PTEN</DIV> Mutational analysis of p53 and PTEN were = performed in=20 the DNA from both tumors and corresponding leukocytes. After amplifying = all=20 exons and intron-exon boundaries, we analyzed the PCR products on 1% = agarose gel=20 and excised and eluted them in sterile H2O. Subsequently, = they were=20 subjected to cycle sequencing reactions using the Thermo = Sequenase=AE=20 Fluorescent Cycle Sequencing kit (Amersham Pharmacia Biotech). The cycle = sequencing products were resolved using a denaturing 6.5% Long Ranger=20 polyacrylamide gel on a sequencing device. PTEN primer sequences = and PCR=20 conditions were previously described [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR12">12</A></CITE>].=20 P53 primer sequences and PCR conditions are based on genome = database=20 entries (<A=20 href=3D"http://www.ncbi.nlm.nih.gov/">http://www.ncbi.nlm.nih.gov/</A>). = Briefly,=20 PCR reactions were carried out using a mixture containing 150 ng = DNA,=20 10 mM Tris, 2 mM MgCl2, HCl (pH 8.3), 0.2 mM = dNTP,=20 10 mM concentrations of each primer and 0.5 unit of Taq DNA = Polymerase.=20 Temperature cycles and times for PCR reactions were: denaturation at = 94=B0C for=20 30 s, annealing at 65=B0C for 30 s, and extension at 72=B0C = for 30 s.=20 Each PCR reaction was preceded by a 3-min denaturation at 94=B0C, and = the final=20 cycle was followed by a 3-min extension at 72=B0C. The total number of = cycles for=20 PCR amplification was 25=9630, depending on the sample DNA. </DIV> <DIV class=3D""><A name=3DSec7></A> <DIV class=3DHeading3>Statistical analysis</DIV> Student t-test, Chi-square test, Fisher=92s exact = test and=20 Mann=96Whitney U-test, were performed to compare differences between = groups=20 depending on the analyzed variables. Spearman correlation test was used = to=20 evaluate the correlation between two parameters. Confidence interval = (CI) was=20 obtained through logistic regression. Survival studies were performed = with the=20 Kaplan Meier analysis and the log-rank test. A value of=20 P < 0.05 was considered to be significant. Analyses = were=20 performed with the SPSS software (version 10, SPSS Inc., Chicago, IL, = USA).=20 </DIV></DIV> <DIV class=3D""><A name=3DSec8></A> <HR> <DIV class=3Dheading2>Results</DIV> <DIV class=3D""><A name=3DSec9></A> <DIV class=3DHeading3>Clinico-pathological characteristics of the = patients</DIV> The male/female ratio was 1:1. The median age at diagnosis = of=20 primary tumor was 59.5 years (range: 27=9669; SD: 12.25). The = median time to=20 tumor recurrence was 7.5 months (range: 3=9624 months; SD: = 5.1; 95%-CI:=20 5.3=967.4 months). With one only exception, all patients displayed = at=20 diagnosis a Karnofsky performance score (KPS) of >70 points. An = association=20 between age or KPS and TTR was not evidenced = (P =3D 0.35,=20 ρ =3D 0.30, Spearman correlation test). There were no = significant=20 associations between mutation of p53 or EGFR amplification = and=20 either survival time (P =3D 0.250 and 0.127, = respectively,=20 log-rank test and Kaplan=96Meier) or time to tumor relapse=20 (P =3D 0.210 and 0.287, respectively, log-rank test and = Kaplan=96Meier). </DIV> <DIV class=3D""><A name=3DSec10></A> <DIV class=3DHeading3>Genetic analyses</DIV> <DIV class=3DPara> <DIV class=3D"">In primary GBM we have observed EGFR = amplification without=20 mutations of p53 in 12 cases, thus indicating a type II GBM = profile. Four=20 further cases showed p53 mutations (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab1">1</A>,=20 Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig1">1</A>)=20 in the absence of EGFR amplification, which was indicative of a = type 1=20 GBM signature. In four GBM neither p53 mutations nor EGFR=20 amplification were identified (non-type 1=96non-type 2 GBM).<A = name=3DTab1></A> <DIV class=3DCapt>Table 1 Sequence = variations=20 of p53 in type 1 glioblastomas </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH align=3Dleft colSpan=3D6> p53 sequence variations </TH></TR> <TR class=3Dheader> <TH align=3Dleft> Case</TH> <TH align=3Dleft> Sequence variation</TH> <TH align=3Dleft> Base exchange</TH> <TH align=3Dleft> Aminoacid substitution</TH> <TH align=3Dleft> CpG site</TH> <TH align=3Dleft> Distribution</TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> G3p</TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.266G>T</TD> <TD align=3Dleft> GGA→GTA</TD> <TD align=3Dleft> Gly→Val</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 44 tumors No germline mutationsa </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G3r</TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.266G>T</TD> <TD align=3Dleft> GGA→GTA</TD> <TD align=3Dleft> Gly→Val</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 44 tumors No germline mutationsa </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G7p</TD> <TD align=3Dleft> c.72C>G (Arg72Pro)</TD> <TD align=3Dleft> CCC→CGC</TD> <TD align=3Dleft> Arg→Pro</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Validated polymorphism Higher risk lung cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR34">34</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS3+21_37dup (P53PIN3) </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Higher risk of CRC [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR36">36</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G7r</TD> <TD align=3Dleft> c.72C>G (Arg72Pro)</TD> <TD align=3Dleft> CCC→CGC</TD> <TD align=3Dleft> Arg→Pro</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Validated polymorphism Higher risk lung cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR34">34</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS3+21_37dup (P53PIN3) </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphismaHigher risk of CRC = [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR36">36</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.278C>T</TD> <TD align=3Dleft> CCT→TCT</TD> <TD align=3Dleft> Pro→Ser</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Somatic mutations in 74 tumors Germline mut. in 2 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G14p</TD> <TD align=3Dleft> c.14G>C</TD> <TD align=3Dleft> CTG→CTC</TD> <TD align=3Dleft> Leu→Leu</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G14r</TD> <TD align=3Dleft> c.14G>C</TD> <TD align=3Dleft> CTG→CTC</TD> <TD align=3Dleft> Leu→Leu</TD> <TD align=3Dleft> No</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G16p</TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS2+38G>C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR41">41</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G16r</TD> <TD align=3Dleft> c.273G>A</TD> <TD align=3Dleft> CGT→CAT</TD> <TD align=3Dleft> Arg→His</TD> <TD align=3Dleft> Yes</TD> <TD align=3Dleft> Somatic mutations in 733 tumors Germline mut. in 16 Li-Fraumenia = </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS2+38G>C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR41">41</A></CITE>]=20 </TD></TR></TBODY></TABLE> <DIV class=3DCapt> <DIV class=3DCaptCont> <DIV class=3D"">aAccording to p53 data bases entries, = mut=20 mutation, CRC colorectal cancer </DIV></DIV></DIV> <DIV class=3DFigure><A name=3DFig1></A><IMG=20 alt=3DMediaObjects/11060_2009_9967_Fig1_HTML.gif=20 src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Fig1_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 1 Upper = row:=20 wild type sequence of p53. Lower row: base exchange = mutation=20 (GGA→GTA) of p53 (exon 8, codon 266), predicting the = aminoacid=20 substitution Gly→Val. The arrow indicates the base exchange </DIV> <HR> </DIV></DIV> <DIV class=3DPara> <DIV class=3D"">At the time of recurrence, every type 1 GBM and type 2 = GBM=20 conserved their genetic pattern concerning p53 mutational status = and=20 EGFR amplification. On the other hand, relapses of non-type = 1=96non-type 2=20 GBM showed in all cases a new EGFR amplification, thus acquiring = the=20 signature of type 2 GBM. Additional PTEN mutations in relapse = cases were=20 observed only among type 2 GBM (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab2">2</A>).=20 Overall, additional LOH were observed more frequently in relapses of = type 2 GBM=20 (83.3%), than in those of type 1 (50%) or non-type 1=96non-type 2 tumors = (25%).=20 Further results are shown in Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>.<A=20 name=3DTab2></A> <DIV class=3DCapt>Table 2 Sequence = variations=20 of PTEN in first and relapse glioblastomas </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <THEAD> <TR class=3Dheader> <TH align=3Dleft colSpan=3D5> PTEN sequence variations </TH></TR> <TR class=3Dheader> <TH align=3Dleft> Case</TH> <TH align=3Dleft> Sequence variation</TH> <TH align=3Dleft> Mutation effect</TH> <TH align=3Dleft> Aminoacid substitution</TH> <TH align=3Dleft> Distribution</TH></TR></THEAD> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft> G1p</TD> <TD align=3Dleft> c.44C>T</TD> <TD align=3Dleft> GGC→GGT</TD> <TD align=3Dleft> Gly→Gly</TD> <TD align=3Dleft> Bladder cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR39">39</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G1r</TD> <TD align=3Dleft> c.44C>T</TD> <TD align=3Dleft> GGC→GGT</TD> <TD align=3Dleft> Gly→Gly</TD> <TD align=3Dleft> Bladder cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR39">39</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G2p</TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G2r</TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G4</TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G4r</TD> <TD align=3Dleft> c.105G>T</TD> <TD align=3Dleft> TGT→TTT</TD> <TD align=3Dleft> Cys→Phe</TD> <TD align=3Dleft> Gliomas [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR38">38</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G5p</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS2-96A>G</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G5r</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.88A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Head neck squamous cell cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR42">42</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G11p</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G11r</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.88A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Head & neck squamous cell cancer [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR42">42</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8-2A>G</TD> <TD align=3Dleft> Splice mutation</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G13p</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G13r</TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS8-2A>G</TD> <TD align=3Dleft> Splice mutation</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.75C>A</TD> <TD align=3Dleft> CAT→AAT</TD> <TD align=3Dleft> His→Asn</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G17p</TD> <TD align=3Dleft> IVS4 109_110 ins TCTTA</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G17r</TD> <TD align=3Dleft> IVS4 109_110 ins TCTTA</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.177A>G</TD> <TD align=3Dleft> TAT→TGT</TD> <TD align=3Dleft> Tyr→Cys</TD> <TD align=3Dleft> Not described mutation</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> IVS5-28_29ins C</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Not described polymorphism</TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G19p</TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> G19r</TD> <TD align=3Dleft> IVS8+32G>T</TD> <TD align=3Dleft> </TD> <TD align=3Dleft> </TD> <TD align=3Dleft> Validated polymorphism Cowden/Bannayan-Riley syndromes [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR40">40</A></CITE>]=20 </TD></TR> <TR class=3Dnoclass> <TD align=3Dleft> </TD> <TD align=3Dleft> c.252A>G</TD> <TD align=3Dleft> GAT→GGT</TD> <TD align=3Dleft> Asp→Gly</TD> <TD align=3Dleft> Glioma [<CITE><A=20 = href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR43">43</A></CITE>]=20 </TD></TR></TBODY></TABLE><A name=3DTab3></A> <DIV class=3DCapt>Table 3 Review = of genetic=20 changes of primary and relapsed glioblastomas </DIV> <TABLE border=3D1> <COLGROUP> <COL align=3Dleft> <COL align=3Dleft></COLGROUP> <TBODY> <TR class=3Dnoclass> <TD align=3Dleft colSpan=3D2> <IMG = alt=3DMediaObjects/11060_2009_9967_Taba_HTML.gif=20 = src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Taba_HTML.gif"=20 align=3Dmiddle> </TD></TR></TBODY></TABLE> <DIV class=3DCapt> <DIV class=3DCaptCont> <DIV class=3D"">TTR time to tumor relapse (months), Mut = mutation,=20 Ampl amplification, p primary (first tumor), r = relapse=20 (first relapse), m male, f female, no ther no = adjuvant=20 therapy performed, RT radiotherapy, A ACNU (nimustine), = V=20 VM26 (teniposide), T temozolomide. Black boxes loss of=20 heterozygosity, grey boxes retention of heterozygosity, white=20 boxes not informative marker </DIV></DIV> <DIV class=3DCaptCont> <DIV class=3D"">* Age (years) at first=20 operation</DIV></DIV></DIV></DIV></DIV> Among p53 mutations there were one C:G to A:T = transition=20 mutation at CpG site (codon 273), one C>T transition at non-CpG site = (codon=20 278), two transversion mutations at non-CpG sites (G>C at codon 14 = and G>T=20 at codon 266, Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig1">1</A>)=20 as well as one validated polymorphism in exon 4 (G>C at codon 72). = Moreover,=20 we have observed two validated intronic polymorphisms in intron 3=20 (IVS3+21_37dup) and intron 2 (IVS2+38G>C). One of the mutations = (c.14G>C)=20 has not been previously described in p53 data bases (<A=20 href=3D"http://http//www.p53.iarc.fr">http://http//www.p53.iarc.fr</A>, = <A=20 href=3D"http://http//www.p53.free.fr">http://http//www.p53.free.fr</A>). = A=20 detailed characterization of p53 sequence variations is shown in=20 Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab1">1</A>.=20 <DIV class=3DPara> <DIV class=3D"">Concerning PTEN mutations, we have identified = four=20 transition mutations in exons 2, 5, 6 and 7, in all cases predicting = aminoacid=20 substitution. Furthermore, there was an acceptor site splice mutation at = the=20 conserved junction nucleotides (IVS8-2A>G). Finally, we have observed = two=20 transversion mutations in exon 5 (c.105G>T) and exon 4 (c.75C>A,=20 Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig2">2</A>)=20 as well as four intronic polymorphisms (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab2">2</A>).=20 The transition in exon 6 (c.177A>G), the splicing mutation and two of = the=20 polymorphisms have not been previously reported (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab2">2</A>).=20 <DIV class=3DFigure><A name=3DFig2></A><IMG=20 alt=3DMediaObjects/11060_2009_9967_Fig2_HTML.gif=20 src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Fig2_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 2 Upper = row:=20 wild type sequence of PTEN. Lower row: base exchange = mutation=20 (CAT→AAT) of PTEN (exon 4, codon 75) predicting the = aminoacid=20 substitution His→Asn. The arrow indicates the base exchange </DIV> <HR> </DIV></DIV> The results of EGFR amplification analysis and = wide-genome=20 LOH are shown in Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>=20 and Supplementary Table 1 (EGFR/CFTR ratios). Interestingly, = GBM=20 cases harboring mutations of p53 always displayed LOH at 17p13 as = well.=20 Similarly, tumors featuring PTEN mutations showed LOH at 10q23, = at least=20 in one of the microsatellites used to analyze this region. = </DIV></DIV> <DIV class=3D""><A name=3DSec11></A> <HR> <DIV class=3Dheading2>Discussion</DIV> GBM mostly exceed in its occurrence and mortality beyond = any other=20 primary brain tumor in adults [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR13">13</A></CITE>].=20 On molecular level, two subsets of GBM have been recognized on the basis = of=20 genetic make-up and clinical features [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR14">14</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR15">15</A></CITE>].=20 Type 2 (de novo) GBM occurs commonly in elderly patients and exhibit=20 overexpression and/or amplification of EGFR. Type 1 (secondary) = GBM shows=20 typically histological progression from a previous low grade astrocytoma = and=20 affects young patients. Paradigmatically, mutations of p53 will = be=20 observed (for review see [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR15">15</A></CITE>]).=20 Most of GBM cases belong to the type 2 group and genetic events are = mutually=20 exclusive for EGFR and p53. Moreover, there are GBM = without either=20 EGFR amplification or p53 mutations (non-type 1=96non-type = 2 GBM)=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR16">16</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR17">17</A></CITE>].=20 During the past decade, accumulated evidence pointed to that type 1 and = type 2=20 GBM constitute distinct disease entities developing through different = genetic=20 pathways [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR13">13</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR14">14</A></CITE>],=20 showing different RNA and protein patterns [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR18">18</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR19">19</A></CITE>]=20 and also probably differing in response to treatment [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR15">15</A></CITE>].=20 After multimodal therapy, GBM almost always recur. = Although this is=20 a capital feature of GBM, only few investigations with series of match = paired=20 patients are available from the literature and available data is = partially=20 contradictory [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR7">7</A></CITE>=96<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR9">9</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR20">20</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR21">21</A></CITE>].=20 We aimed to investigate whether molecular subtypes of GBM also show = different=20 profiles at the time of relapse. <DIV class=3DPara> <DIV class=3D"">In our genetic analysis, we have observed two patterns = of tumor=20 recurrence (Fig. <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Fig3">3</A>):=20 GBM type 1 and type 2 retained upon recurrence their genetic alterations = affecting p53 and EGFR. In contrast, all relapses of = non-type=20 1=96non-type 2 GBM showed a new EGFR amplification, thus they = acquired a=20 type 2 GBM profile. Accumulation of additional genetic alterations upon = relapse=20 included PTEN mutations in four type 2 GBM (but not in the other=20 subtypes). Additional LOH were more frequently observed in relapses of = type 2=20 GBM (in 10/12 cases) compared to relapses of type 1 GBM and non-type = 1=96non-type=20 2 GBM (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>).=20 The accumulation of alterations during the clonal expansion of = transformed glial=20 cells is well known in GBM, confer tumor cells a growth advantage and = underlies=20 the genetic heterogeneity of GBM [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR22">22</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR23">23</A></CITE>].=20 This phenomenon contributes to explain the molecular differences between = primary=20 tumors and relapses [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR9">9</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR24">24</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR25">25</A></CITE>].=20 <DIV class=3DFigure><A name=3DFig3></A><IMG=20 alt=3DMediaObjects/11060_2009_9967_Fig3_HTML.gif=20 src=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects/= 11060_2009_9967_Fig3_HTML.gif"></DIV> <DIV class=3DCapt>Fig. 3 Graphic = showing two=20 molecular patterns of GBM recurrence </DIV> <HR> </DIV></DIV> As a part of one previous investigation we had = characterized the=20 promoter methylation status of the DNA repair gene MGMT in all = type 1 GBM=20 (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>,=20 cases G3, G7, G14 and G16), non-type 1=96non-type 2 GBM (Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab3">3</A>,=20 cases G2, G6, G8 and G9) as well as eight type 2 GBM here described = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR26">26</A></CITE>].=20 This analysis together with the present results of p53 and = EGFR=20 have evidenced similar rates of MGMT methylation for the three = GBM=20 subtypes, and they are in accordance with the overall methylation rates = reported=20 in glioblastoma [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR5">5</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR27">27</A></CITE>].=20 Typically, MGMT was methylated in the first GBM and remained = always=20 methylated upon relapse. Thus, our results also confirmed the = observation that,=20 in the pathogenesis and progression of GBM, MGMT becomes = hypermethylated=20 at an early stage [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR28">28</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR29">29</A></CITE>]=20 and remains methylated through progression, a situation which was has = been=20 described in small adenomas and carcinomas of colorectal origin as well=20 [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR30">30</A></CITE>].=20 The p53 mutational analysis in those type 1 GBM with methylated=20 MGMT showed G:C to A:T transition within CpG site (c.273 = CGT→CAT) in two=20 cases. Further mutations of p53 in tumors with methylated = MGMT (1=20 case) occurred in non-CpG sites. From p53 data bases entries,=20 approximately 52% of mutational events are transition mutations and, of = this=20 subset, about 72% are G:C to A:T transitions (<A=20 href=3D"http://http//www.p53.iarc.fr">http://http//www.p53.iarc.fr</A>, = <A=20 href=3D"http://http//www.p53.free.fr">http://http//www.p53.free.fr</A>) = [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR31">31</A></CITE>],=20 although the signature of the mutational spectrum varies according to = tumor=20 type. It was previously suggested a link [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR29">29</A></CITE>]=20 between MGMT methylation and the presence of G:C to A:T = transition=20 mutations in p53 in gliomas. In the present study we could = confirm the=20 former in two patients (cases G14p, G14r, G16p and G16r, Table <A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#Tab1">1</A>);=20 in these cases, G:C to A:T transitions could be attributable to a = inactivation=20 in MGMT that allows O 6 -methylguanine = to=20 persist and be read as an adenine. Our observations contrast with a = recent=20 investigation about GBM reporting only 4% of mutations of p53 in = tumors=20 with methylated MGMT within CpG sites [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR32">32</A></CITE>]=20 although we should take in mind the size of our type 1 GBM collective, = clearly=20 too short to state a definite tendency. Strikingly, we have identified in cases G7p and G7r the = exonic=20 p53 polymorphism c.72C>G (Arg72Pro) together with the intronic = polymorphism IVS3+21_37dup (P53PIN3). Several investigations have = pointed=20 the association between p53 polymorphisms and an increased risk = for=20 different cancers: for instance, in the case of the exonic c.72 variant = such=20 association was suggested since the p53 protein with Arg72 is more = efficient in=20 inducing apoptosis than the one with the Pro72 variant [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR33">33</A></CITE>].=20 In clinical studies, the variants c.72 Pro/Pro and c.72 Arg/Pro have = been=20 associated with an elevated risk of lung cancer [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR34">34</A></CITE>,=20 <CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR35">35</A></CITE>].=20 Recently, intron 3 duplication (IVS3+21_37dup) has been found to be = associated=20 with increased risk of colorectal cancer [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR36">36</A></CITE>].=20 However, the authors could not determine whether this intron 3 = duplication alone=20 influences mRNA stability or if this effect requires the codon Pro72 = variant. To=20 our knowledge, the occurrence of both polymorphisms in GBM patients has = not been=20 reported before and pointed to a possible role in brain cancer as well. = With respect to PTEN, we have identified mutations = only in=20 type 2 GBM. In four cases, PTEN mutations occurred in both the = first GBM=20 and the relapse. In four further cases, new mutations were observed in = the=20 recurrences but not in the first tumor. The rate and pattern of = mutations are in=20 accordance with previous reported data [<CITE><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/fulltext.htm= l#CR37">37</A></CITE>].=20 All tumors with PTEN mutations also harbored LOH at the = chromosome region=20 10q23, which pointed to the inactivation of this tumor suppressor gene = and make=20 patent its contribution to the tumorigenesis and progression in GBM. = Taken together, our results strongly suggest that GBM = relapses,=20 compared to the corresponding first tumor, accumulate additional = molecular=20 alterations and may develop along two distinct pathways, which = contribute to=20 delineate the different profiles of type 1, type 2 and non-type = 1=96non-type 2=20 GBM. </DIV> <DIV class=3DAcknowledgments>Open=20 Access This article is distributed = under the=20 terms of the Creative Commons Attribution Noncommercial License which = permits=20 any noncommercial use, distribution, and reproduction in any medium, = provided=20 the original author(s) and source are credited.</DIV> <DIV class=3DAppendix><A name=3DAppESM1></A> <DIV class=3D""><A name=3DSecESM1></A> <HR> <DIV class=3Dheading2>Electronic supplementary material</DIV> <DIV class=3DPara> <DIV class=3D"">Below is the link to the electronic supplementary = material.=20 <DIV><A=20 href=3D"http://www.springerlink.com/content/j12217q235065261/MediaObjects= /11060_2009_9967_MOESM1_ESM.pdf"> <DIV class=3DCapt>Supplementary material 1 (PDF 25=20 kb)</DIV></A></DIV></DIV></DIV></DIV></DIV> <HR> <H2><A name=3DBib1></A>References </H2> <TABLE> <TBODY class=3DCitation> <TR vAlign=3Dtop> <TD>1.</TD> <TD><A name=3DCR1></A>Salcman M, Kaplan R (1991) Intracranial tumors = in=20 adults. 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