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{=0A= document.writeln('

3D"Click

');=0A= }=0A= =0A= function get_text_link(url, linktext)=0A= {=0A= document.writeln('' + = linktext + '');=0A= }=0A= =0A= function get_htmlftext_link(a, b) {=0A= document.writeln("" + = b + ", ");=0A= }=0A= =0A= ------=_NextPart_000_0005_01CA352F.3D8DF910 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://download.interscience.wiley.com/freeflow/js/personalisation.js /*=0A= * Project : Interscience Redesign=0A= * File : personalisation.js=0A= * Creation Date : 25th March 2003=0A= * Author : Jonathan Sugar=0A= *=0A= * Various functions to deliver personalisation functionality.=0A= */=0A= =0A= // Pre-declare userName so it's available when we need to test if the = page=0A= // assembler ran OK=0A= var userName =3D null;=0A= var shcrtCookie =3D null;=0A= =0A= // Browser detect globals:=0A= var uaStr =3D navigator.userAgent.toLowerCase();=0A= var uaVer =3D parseInt(navigator.appVersion);=0A= // for detecting netscape 4=0A= var netscape =3D (uaStr.indexOf("mozilla") > -1);=0A= var netscape4 =3D (netscape && uaVer < 5);=0A= =0A= // Factory Methods=0A= =0A= // Simple wraper object to encapsulate the data for a co-branding item.=0A= function coBrandingItem(image, imageAltText, href)=0A= {=0A= //alert("Making new coBrandingItem:\nImage: " + image + = "\nimageAltText" + imageAltText + "\nhref: " + href);=0A= this.image =3D image;=0A= this.imageAltText =3D imageAltText;=0A= this.href =3D href;=0A= }=0A= =0A= /*=0A= * Insert the 'Customer co-branding' section.=0A= * -----------------------------=0A= *=0A= * Author: Jonathan Sugar, March 2003.=0A= *=0A= * 1. Insert customer co branding. Loop through 'customerCoBranding' array=0A= * inserting XHTML where needed.=0A= *=0A= * NOTE: the array contains ONLY the resource location (the location of a = gif=0A= * on the file system), the required XHTML is then added where needed. = This way=0A= * we maintain the seperation between presentation and content.=0A= */=0A= function insertCustomerCoBranding(maxBranding)=0A= {=0A= if( customerCoBranding[0].image !=3D "none")=0A= {=0A= document.writeln('

');=0A= document.writeln('

');=0A= document.writeln('

' + imageAltText + '
' + imageAltText + = '
More Providers

');=0A= document.writeln('

');=0A= document.writeln('

3D""

');=0A= }=0A= }=0A= =0A= /*=0A= * Insert the 'My Area' section.=0A= * -----------------------------=0A= *=0A= * Author: Jonathan Sugar, March 2003.=0A= * Updated: Ben Marvell, April 2006=0A= *=0A= * My profile link absolute as this file isnt only run from w3 anymore.=0A= */=0A= function insertMyArea(userdata)=0A= {=0A= // just in case the page assembler isn't running.=0A= =0A= =0A= if (userName !=3D null)=0A= {=0A= document.writeln("

" + = userName.substring(0,33) + "..., access your
" + userName + = ", access your

My Profile"); =0A= } =0A= document.writeln(getLoginOutLink());=0A= //CR 3652 getAthenStatus() as added=0A= document.writeln(getAthenStatus());=0A= document.writeln("

");=0A= } else {=0A= if(typeof(environment)!=3D'undefined' && = environment=3D=3D'PRODUCTION') {=0A= document.writeln("

Log Out");=0A= document.writeln("

"); =0A= }=0A= }=0A= if (menuloadedflag){=0A= wisMenu();=0A= }=0A= }=0A= =0A= /*=0A= * Insert the Customer Admin 'Login/Logout' link=0A= * -----------------------------=0A= *=0A= * Author: Bobby Jack, December 2004.=0A= *=0A= */=0A= function insertAdminLink(page)=0A= {=0A= val =3D getCookie("AID");=0A= =0A= if (val && val !=3D "-1")=0A= {=0A= document.writeln('

Logout');=0A= }=0A= else=0A= {=0A= document.writeln('

');=0A= }=0A= }=0A= =0A= /*=0A= * Insert the username section.=0A= * -----------------------------=0A= *=0A= * Author: Jonathan Sugar, March 2003.=0A= *=0A= * 1. Insert the username.=0A= */=0A= function insertUserName()=0A= {=0A= document.write(userName);=0A= }=0A= =0A= /*=0A= * Return the appropriate link to the login/logout cgi=0A= * ---------------------------------------------------=0A= *=0A= * Author: Jonathan Sugar, March 2003.=0A= * Updated: Ben Marvell, April 2006=0A= * Had to change the links absolute as this file isnt only run from w3 = anymore.=0A= */=0A= function getLoginOutLink()=0A= {=0A= if(userName !=3D "")=0A= {=0A= return "

";=0A= }else=0A= {=0A= return "

";=0A= }=0A= }=0A= =0A= /*=0A= * Return the appropriate link to the shopping cart content page=0A= * ---------------------------------------------------=0A= *=0A= * Author: Zhiming & Warrell June 2007=0A= */=0A= function getShCrtLink(logged)=0A= {=0A= if(logged =3D=3D true )=0A= return "

";=0A= else=0A= return "

";=0A= }=0A= =0A= function toggleShCrtLink()=0A= {=0A= shcrtCookie =3D readCookie("SHCRT");=0A= if(shcrtCookie !=3D null)=0A= {=0A= if(document.all)=0A= {=0A= return true;=0A= }=0A= else=0A= {=0A= document.getElementById("shcrt_id").style.display =3D "block";=0A= }=0A= }=0A= else=0A= {=0A= if(document.all)=0A= {=0A= return true;=0A= }=0A= else=0A= {=0A= document.getElementById("shcrt_id").style.display =3D "none";=0A= }=0A= }=0A= }=0A= =0A= =0A= =0A= function readCookie(name)=0A= {=0A= var nameEQ =3D name + "=3D";=0A= var ca =3D document.cookie.split(';');=0A= for(var i=3D0; iYou are logged in = to Athens';=0A= }=0A= else=0A= {=0A= return "

";=0A= }=0A= =0A= } ------=_NextPart_000_0005_01CA352F.3D8DF910 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://download.interscience.wiley.com/freeflow/js/site.js /************************************************************************= ***************************=0A= FILE: site.js=0A= AUTHOR: Jonathan Sugar, Chris Neatby-Smith and Digital Pulp. July = 2003.=0A= =0A= DESCRIPTION: Javascript relating to site wide functionality across =0A= InterScience.=0A= =0A= INCLUDES FUNCTIONS=0A= =3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=0A= =0A= Search Functions=0A= ----------------=0A= =0A= selectAndSubmit() - for deciding where to submit a query based on the = state of the form.=0A= processallContentSearch() - for validating form input for an all = content search.=0A= processTitleSearch() - for validating form input for a Title search.=0A= checkCitationSearch() - for validating the citation search form.=0A= selectAll() - check all the checkboxes in a form.=0A= do_popup() - stasa Acronym Finder window popup function.=0A= =0A= =0A= Other Functions=0A= ---------------=0A= =0A= fixInlineLists() - fix netscape 4.x bug where

tags with images = display both an image and=0A= the default bullet. =0A= =0A= = *************************************************************************= **************************/=0A= =0A= var http =3D false;=0A= if(navigator.appName =3D=3D "Microsoft Internet Explorer") {=0A= http =3D new ActiveXObject("Microsoft.XMLHTTP");=0A= } else {=0A= http =3D new XMLHttpRequest();=0A= } =0A= =0A= // Global Variables=0A= var debug =3D false;=0A= var first =3D true;=0A= =0A= // Browser detect globals:=0A= var uaStr =3D navigator.userAgent.toLowerCase();=0A= var uaVer =3D parseInt(navigator.appVersion);=0A= // for detecting netscape 4=0A= var netscape =3D (uaStr.indexOf("mozilla") > -1);=0A= var netscape4 =3D (netscape && uaVer < 5);=0A= =0A= /*=0A= * Used to switch search mode between APS and TitleFinder on quicksearch = boxes=0A= * = -------------------------------------------------------------------------= --=0A= *=0A= * 1. Decides which search is required based on the value of the radio = buttons=0A= * in 'document.formSelector.whichForm'.=0A= *=0A= * 2. Calls individual functions to process the forms and submit them.=0A= *=0A= */=0A= function selectAndSubmit()=0A= {=0A= var formSelection;=0A= =0A= // find out which form we need to process.=0A= var selector =3D document.formSelector.whichForm;=0A= for (var i =3D 0; i < selector.length; i++){=0A= if (selector[i].checked){=0A= formSelection =3D selector[i].value;=0A= }=0A= }=0A= =0A= // Call the relevent function to process the required form.=0A= if(formSelection =3D=3D "allContentSearch"){=0A= processallContentSearch(document.allContentSearchForm, = document.formSelector)=0A= }=0A= else if(formSelection =3D=3D "searchTitles"){=0A= processTitleSearch(document.searchTitles, document.formSelector)=0A= }=0A= else{=0A= return false;=0A= }=0A= =0A= //@@@DM It was anoying me this error =0A= return false;=0A= }=0A= =0A= /*=0A= *Process the ADS search form=0A= *=0A= */=0A= function trim(str)=0A= {=0A= return str.replace(/(^\s*)|(\s*$)/g,"");=0A= }=0A= =0A= function ADSquery(t) {=0A= var keyword =3D trim(t.getElementsByTagName("input")[0].value);=0A= if (keyword!=3D"") {=0A= = window.open("","popup","toolbar=3Dyes,location=3Dyes,status=3Dyes,menubar= =3Dno,scrollbars=3Dyes,resizable=3Dyes,width=3D800,height=3D500");=0A= t.target=3D"popup";=0A= t.qsearch.value=3Dkeyword;=0A= return true;=0A= = //processADSSearch(document.ADSSearchForm,document.ADSQueryForm,keyword);=0A= } else {=0A= alert("Please input a keyword for ADS searching before clicking go");=0A= return false;=0A= }=0A= }=0A= /*=0A= function processADSSearch(hiddenForm ,queryForm , str) {=0A= var terms =3D str;=0A= var formToSubmit =3D hiddenForm;=0A= var formToProcess =3D queryForm;=0A= formToSubmit.qsearch.value =3D terms;=0A= = window.open("","popup","toolbar=3Dyes,location=3Dyes,status=3Dyes,menubar= =3Dno,scrollbars=3Dyes,resizable=3Dyes,width=3D800,height=3D500");=0A= formToSubmit.action =3D = "http://adsabs.harvard.edu/cgi-bin/nph-basic_connect"=0A= formToSubmit.target =3D "popup";=0A= formToSubmit.submit();=0A= }=0A= */=0A= =0A= /*=0A= * Process the allContentSearch form=0A= * ---------------------------------=0A= * 1. Perform form validation, checking for empty fields.=0A= * Give the user a relevent message if required.=0A= *=0A= * 2. Process/Assign the appropriate form elements and submit the form.=0A= */=0A= function processallContentSearch(hiddenForm, queryForm)=0A= {=0A= var formToSubmit =3D hiddenForm;=0A= var formToProcess =3D queryForm;=0A= =0A= // Do some basic form validation=0A= if(formToProcess.terms.value !=3D "")=0A= {=0A= formToSubmit.WISsearch1.value =3D formToProcess.terms.value;=0A= formToSubmit.submit(); =0A= }=0A= else=0A= {=0A= alert("You must enter a search term in the text box before you click = go.");=0A= }=0A= }=0A= =0A= /*=0A= * Process the Title Search form=0A= * -----------------------------=0A= * 1. Perform form validation, checking for empty fields.=0A= * Give the user a relevent message if required.=0A= *=0A= * 2. Process/Assign the appropriate form elements and submit the form.=0A= */=0A= =0A= function processTitleSearch(hiddenForm, queryForm)=0A= {=0A= var formToSubmit =3D hiddenForm;=0A= var formToProcess =3D queryForm;=0A= =0A= // Do some basic form validation=0A= if(formToProcess.terms.value !=3D "")=0A= {=0A= formToSubmit.query.value =3D formToProcess.terms.value;=0A= formToSubmit.submit();=0A= }=0A= else=0A= {=0A= alert("You must enter a search term in the text box before you click = go.");=0A= }=0A= }=0A= =0A= function doCitationSearch()=0A= {=0A= var form =3D document.citsearchform;=0A= if(checkCitationSearch())=0A= {=0A= form.action =3D "/search/allsearch";=0A= form.submit();=0A= }=0A= }=0A= =0A= // Preprocesses the arguments sent to CitationSearch and checks validity = of user inputs.=0A= function checkCitationSearch()=0A= {=0A= var supplementFound =3D 0; //increment if found;=0A= var hyphenFound =3D 0; //increment if found;=0A= var fm =3D document.citsearchform;=0A= var errmsg =3D "Please use the valid digits '0' to '9'";=0A= // remove all spaces=0A= fm.volume.value =3D fm.volume.value.replace(/ /g, "");=0A= fm.issue.value =3D fm.issue.value.replace(/ /g, "");=0A= fm.pages.value =3D fm.pages.value.replace(/ /g, "");=0A= =0A= if (fm.issn.value.length > 0 && fm.volume.value.length > 0 && = fm.pages.value.length > 0)=0A= {=0A= // check page range=0A= var pageRange =3D fm.pages.value;=0A= var volume =3D fm.volume.value;=0A= var issue =3D fm.issue.value;=0A= var continueFlag =3D 0;=0A= //check the volume=0A= // if (isNaN(volume))=0A= // {=0A= // // volume failed checking=0A= // alert("Volume is invalid. "+errmsg+".");=0A= // return false;=0A= // }=0A= //check the issue=0A= if ((issue.length !=3D 0) && isNaN(issue))=0A= {=0A= // issue failed checking=0A= alert("Issue is invalid. "+errmsg+" or leave blank.");=0A= return false;=0A= }=0A= //check the page range=0A= if (isNaN(pageRange))=0A= {=0A= var isValid =3D false;=0A= // check for hyphen=0A= var hyphenIndex =3D pageRange.indexOf("-");=0A= if ((hyphenIndex !=3D -1) && (hyphenIndex =3D=3D = pageRange.lastIndexOf("-")))=0A= {=0A= // we have only one hyphen, so extract each number=0A= var firstNum =3D pageRange.substring(0, hyphenIndex);=0A= var secondNum =3D pageRange.substring(hyphenIndex + 1);=0A= // check for S=0A= var firstS =3D firstNum.indexOf("S");=0A= var secondS =3D secondNum.indexOf("S");=0A= if (firstS =3D=3D secondS)=0A= {=0A= if ((firstS =3D=3D -1) && isNum(firstNum) && isNum(secondNum) && = (firstNum.length > 0) && (secondNum.length > 0))=0A= isValid =3D true; // both inputs are digits=0A= else if (firstS =3D=3D 0)=0A= {=0A= var firstNumNoS =3D firstNum.substring(1);=0A= var secondNumNoS =3D secondNum.substring(1);=0A= if (isNum(firstNumNoS) && isNum(secondNumNoS) && = (firstNumNoS.length > 0) && (secondNumNoS.length > 0))=0A= isValid =3D true; // both inputs are Sdigits=0A= }=0A= =0A= }=0A= }=0A= if (isValid)=0A= {=0A= return true;=0A= }=0A= else=0A= {=0A= alert("The page range is invalid. "+errmsg+" and a hyphen '-' if = required.");=0A= return false;=0A= }=0A= }=0A= return true;=0A= }=0A= else=0A= {=0A= alert("Volume and pages are required fields. Please make sure data has = been entered for both fields.");=0A= return false;=0A= }=0A= }=0A= =0A= // returns true if a is an Integer.=0A= function isNum(a)=0A= { return !(isNaN(a)); }=0A= =0A= function fixInlineLists() {=0A= if (!document.getElementById) return;=0A= var allLIs =3D document.getElementsByTagName('li');=0A= var bulletNode =3D document.createElement('img');=0A= bulletNode.src = =3D"http://download.interscience.wiley.com/images/icon.bullet.gif";=0A= =0A= for (var i=3D0; iAbstract | References';=0A= }=0A= //alert("URL: " + loc + "\nStart Index: " + startIndex + "\nEnd Index: = " + endIndex + "\nPage: " + page + "\nLink Text: " + links);=0A= return links;=0A= }=0A= =0A= =0A= function selectAll(fm)=0A= {=0A= //var fm =3D document.viewSelectedForm;=0A= =0A= for (var i =3D 0; i < fm.elements.length; i++)=0A= {=0A= if (fm.elements[i].type =3D=3D 'checkbox' /*&& fm.elements[i].name = =3D=3D 'ID'*/)=0A= {=0A= fm.elements[i].checked =3D true;=0A= }=0A= }=0A= }=0A= =0A= =0A= function do_popup(file)=0A= {=0A= var w =3D window.open(file, 'popWin', = 'width=3D350,height=3D500,scrollbars=3Dyes');=0A= =0A= }=0A= =0A= =0A= function mailTool(itemID, useurl, skin)=0A= {=0A= //var url =3D "/cgi-bin/mailform?item=3D"+itemID;=0A= var url =3D "/cgi-bin/mailform?item=3D" + itemID + "&skin=3D" + skin;=0A= var parameters =3D = 'status=3Dno,scrollbars=3Dno,resizable=3Dyes,width=3D400,height=3D475';=0A= =0A= if (useurl !=3D "")=0A= {=0A= url +=3D "&useurl=3D"+useurl;=0A= }=0A= =0A= if ((isNaN(parseInt(itemID))=3D=3Dtrue)||(arguments.length >=3D 4 && = arguments[3] =3D=3D 'cluster'))=0A= {=0A= url +=3D '&type=3Dcluster';=0A= }=0A= =0A= //check whether the last 2 arguments is used for urllink.=0A= // urllink is needed to open up mail form for mrw pages on tomcat = port=0A= if (arguments.length>=3D5 && = arguments[arguments.length-2]=3D=3D'urllink' && = arguments[arguments.length-1]!=3D'')=0A= {=0A= url =3D arguments[arguments.length-1] + url;=0A= }=0A= =0A= if (arguments.length>=3D5 && arguments[3]=3D=3D'currentlink' && = arguments[4]!=3D'')=0A= {=0A= url =3D url + '¤tLink=3D' + arguments[4];=0A= =0A= //processing specific cases=0A= if (arguments[4]=3D=3D'jcatalogjsphdr')=0A= {=0A= //get the issn and pass it the next stage for processing=0A= var currloc =3D location.href;=0A= var issnloc =3D currloc.indexOf("&issn=3D");=0A= if (issnloc !=3D -1)=0A= {=0A= url =3D url + '¤tISSN=3D' + = currloc.substring(issnloc+6);=0A= }=0A= }=0A= // issn is needed for cocoon rendered mrw to form the final url.=0A= else if ( (arguments[4]=3D=3D'cocoonmrwhome') || = (arguments[4]=3D=3D'cocoonmrwarticle') )=0A= {=0A= if (arguments.length>=3D7 && arguments[5]=3D=3D'issn' && = arguments[6]!=3D'')=0A= {=0A= url =3D url + '¤tISSN=3D' + arguments[6];=0A= }=0A= //getting the parameters of the currently shown mrw from the = url=0A= //the method used here is a little crude, it extracts the = values from the current url of the page the user is in =0A= //to pass it to the background program to reconstruct the = requred url=0A= if (arguments[4]=3D=3D'cocoonmrwhome')=0A= {=0A= var currloc =3D location.href;=0A= var mrwParamArr =3D currloc.split("/");=0A= if (mrwParamArr.length >=3D 7)=0A= {=0A= url =3D url + '&mrwPath=3D' + mrwParamArr[5];=0A= var mrwFileParam =3D mrwParamArr[6];=0A= var seploc =3D mrwFileParam.indexOf("#");=0A= if (seploc !=3D -1)=0A= {=0A= url =3D url + '&mrwSection=3D' + = mrwFileParam.substring(seploc+1);=0A= mrwFileParam =3D = mrwFileParam.substring(0,seploc);=0A= }=0A= seploc =3D mrwFileParam.indexOf("?");=0A= if (seploc !=3D -1)=0A= {=0A= url =3D url + '&mrwUrlParam=3D' + = mrwFileParam.substring(seploc+1);=0A= url =3D url + '&mrwDocType=3D' + = mrwFileParam.substring(0,seploc);=0A= }=0A= else=0A= {=0A= url =3D url + '&mrwDocType=3D' + mrwFileParam;=0A= }=0A= }=0A= =0A= }=0A= else if (arguments[4]=3D=3D'cocoonmrwarticle')=0A= {=0A= var currloc =3D location.href;=0A= var mrwParamArr =3D currloc.split("/");=0A= if (mrwParamArr.length >=3D 7)=0A= {=0A= url =3D url + '&mrwPath=3D' + mrwParamArr[5];=0A= url =3D url + '&mrwAid=3D' + mrwParamArr[7];=0A= url =3D url + '&mrwVersion=3D' + mrwParamArr[8];=0A= var mrwFileParam =3D mrwParamArr[9];=0A= var seploc =3D mrwFileParam.indexOf("#");=0A= if (seploc !=3D -1)=0A= {=0A= url =3D url + '&mrwSection=3D' + = mrwFileParam.substring(seploc+1);=0A= url =3D url + '&mrwDocType=3D' + = mrwFileParam.substring(0,seploc);=0A= }=0A= else=0A= {=0A= url =3D url + '&mrwDocType=3D' + mrwFileParam;=0A= }=0A= }=0A= }=0A= }=0A= }=0A= =0A= //checking if a html file exists in the url=0A= var tmpfname =3D location.href;=0A= var loc =3D tmpfname.indexOf("?");=0A= if (loc !=3D -1)=0A= {=0A= tmpfname =3D tmpfname.substring(0,loc);=0A= }=0A= loc =3D tmpfname.indexOf(".html");=0A= if (loc !=3D -1)=0A= {=0A= loc =3D tmpfname.lastIndexOf("/");=0A= if (loc !=3D -1)=0A= {=0A= tmpfname =3D tmpfname.substring(loc+1);=0A= url =3D url + '¤tFile=3D' + tmpfname;=0A= }=0A= }=0A= =0A= if (arguments.length>=3D7 && arguments[5]=3D=3D'currentyear' && = arguments[6]!=3D'')=0A= {=0A= url =3D url + '¤tYear=3D' + arguments[6];=0A= }=0A= =0A= eval("wileyMailTool =3D = window.open('"+url+"','MailTool','"+parameters+"')");=0A= =0A= // If object exists ensure has focus=0A= if (eval("wileyMailTool") && window.focus)=0A= eval("wileyMailTool").focus();=0A= }=0A= =0A= =0A= /*=0A= * 1. Find the links we need to alter.=0A= * 2. For each link apply the new styles we need.=0A= */=0A= function fixAbsRefRendering()=0A= { =0A= var table =3D document.getElementById('prerendered');=0A= var anchors =3D table.getElementsByTagName('a');=0A= for (var i =3D 1; i < anchors.length; i++)=0A= {=0A= if(anchors[i].href !=3D"") =0A= { =0A= applyAbsRefStyleCorrection(anchors[i]);=0A= =0A= }=0A= }=0A= }=0A= =0A= function applyAbsRefStyleCorrection(currentLink)=0A= {=0A= currentLink.style["color"] =3D "#369";=0A= currentLink.style["fontWeight"] =3D "normal";=0A= currentLink.style["marginLeft"] =3D "3px";=0A= currentLink.style["marginRight"] =3D "5px";=0A= }=0A= =0A= function populatePage(formName)=0A= {=0A= if (formName.Page.value =3D=3D '')=0A= {=0A= formName.Page.value =3D document.location;=0A= }=0A= }=0A= =0A= function equalizeColumnWidth(numFeatures)=0A= {=0A= var numberOfFeatures =3D numFeatures; =0A= // Percentage width of table to use for each column.=0A= var percentPerFeature =3D Math.floor(100 / numberOfFeatures); =0A= // Number of columns in total. Content cols + seperators. =0A= =0A= document.writeln("");=0A= for(var i =3D 0; i < numberOfFeatures; i++)=0A= { =0A= document.writeln(' 3D"blank"

');=0A= }=0A= document.writeln("");=0A= }=0A= =0A= function old_equalizeColumnWidth(numFeatures)=0A= {=0A= var numberOfFeatures =3D numFeatures; =0A= // Percentage width of table to use for each column.=0A= var percentPerFeature =3D Math.floor(100 / numberOfFeatures); =0A= // Number of columns in total. Content cols + seperators. =0A= =0A= if(debug)=0A= {=0A= alert("Number of feature boxes: " + numberOfFeatures + "\nPercent per = feature: " + percentPerFeature);=0A= }=0A= =0A= document.writeln("");=0A= for(var i =3D 0; i < numberOfFeatures; i++)=0A= { =0A= if (netscape4 =3D=3D false)=0A= {=0A= document.writeln(' 3D"blank"

'); =0A= }=0A= }=0A= document.writeln("");=0A= }=0A= =0A= function writeFBoxCell()=0A= {=0A= if(first)=0A= {=0A= document.write('');=0A= }else=0A= {=0A= document.write('');=0A= }=0A= numFeatures ++;=0A= }=0A= =0A= =0A= //=0A= // escape apostrophe characters within a string=0A= //=0A= function encode(str)=0A= {=0A= out =3D '';=0A= =0A= for (a =3D 0; a < str.length; a++)=0A= {=0A= c =3D str.charAt(a);=0A= =0A= if (c =3D=3D "'")=0A= {=0A= out +=3D '\\';=0A= }=0A= =0A= out +=3D c;=0A= }=0A= =0A= return out;=0A= }=0A= =0A= =0A= //=0A= //=0A= //=0A= function escapeEntities(str)=0A= {=0A= while (str.indexOf("'") > -1)=0A= {=0A= pos =3D str.indexOf("'");=0A= str =3D str.substr(0, pos) + '\\\'' + str.substr(pos + 6);=0A= }=0A= =0A= return str;=0A= }=0A= =0A= =0A= // new popup function less intrusive just uses the DOM to pickup links = with an exwin class.=0A= // currently implemented for Figure popups on D&T homepage.=0A= =0A= function exLinks()=0A= {=0A= if (document.getElementById)=0A= {=0A= var aObject =3D document.getElementsByTagName("a");=0A= for (var x=3D0; x" + = liels[j].innerHTML + "

You do not have = anything saved in this menu.

";=0A= else=0A= tmpinnerhtml +=3D "

" = + liels[j].innerHTML + "

"; =0A= } =0A= } else {=0A= //setup var to see if more is required=0A= var more =3D (subnodeslength < lilength) ? true : = false; =0A= var flag =3D true;=0A= for (var j=3D0; j" + liels[j].innerHTML + = "";=0A= } else if (j > lilength-3){=0A= if (j =3D=3D lilength-2){=0A= var url =3D liels[j].innerHTML.split('"')[1];=0A= tmpinnerhtml +=3D "

" + liels[j].innerHTML + "

";=0A= } else {=0A= tmpinnerhtml +=3D "

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Cancer Science

ISSN: 1349-7006 (Online)

ISSN: 1347-9032 (Print)

Volume 9999, Issue 9999, 2009.

Pages: ???=96???

Top of Page
Abstract
Definition of Glioma Types
EGFR/RAS/NF1/PTEN/PI3K Pathway
TP53/MDM2/MDM4/p14^ARF = Pathway
p16^INK4a/CDK4/RB1 Pathway
IDH1
Loss of Heterozygosity (LOH)
Loss of 1p and 19q
cDNA and Protein Expression Profiles
Response to Chemotherapy
Chemotherapy-induced Genetic Alterations
Brain Tumor Stem Cells
Outlook
References

List of=20 Images

Abstract=20 Only

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3D"About

Online ISSN: = 1096-9896 Print ISSN: 0022-3417
The Journal of = Pathology
Volume 214, Issue 3, 2008. Pages: 328-336

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Review Article =

Genetic alterations and signaling = pathways in=20 the evolution of gliomas

Hiroko Ohgaki=20 ¹ and Paul Kleihues

International Agency = for Research=20 on Cancer (IARC), Lyon, France; Department of Pathology, = University of=20 Z=FCrich, Z=FCrich, Switzerland

¹ To whom = correspondence=20 should be addressed. E-mail: ohgaki@iarc.fr

Copyright =A9 2009 Japanese Cancer=20 Association

ABSTRACT

3DAbstract=20

Gliomas are the most common primary brain tumors. = They=20 account for more than 70% of all neoplasms of the central nervous = system=20 and vary considerably in morphology, location, genetic = alterations, and=20 response to therapy. Most frequent and malignant are = glioblastomas. The=20 vast majority (>90%) develops rapidly after a short clinical = history=20 and without evidence of a less malignant precursor lesion (primary = or=20 de novo glioblastoma). Secondary = glioblastomas=20 develop more slowly through progression from low-grade or = anaplastic=20 astrocytoma. These glioblastoma subtypes constitute distinct = disease=20 entities that affect patients of different age, develop through = distinct=20 genetic pathways, show different RNA and protein expression = profiles, and=20 may differ in their response to radio- and chemotherapy. Recently, = isocitrate dehydrogenase 1 (IDH1) mutations = have been=20 identified as a very early and frequent genetic alteration in the = pathway=20 to secondary glioblastomas as well as that in oligodendroglial = tumors,=20 providing the first evidence that low-grade astrocytomas and=20 oligodendrogliomas may share common cells of origin. In contrast, = primary=20 glioblastomas very rarely contain IDH1 = mutations,=20 suggesting that primary and secondary glioblastomas may originate = from=20 different progenitor cells, despite the fact that they are = histologically=20 largely indistinguishable. In this review, we summarize the = current status=20 of genetic alterations and signaling pathways operative in the = evolution=20 of astrocytic and oligodendroglial tumors. (Cancer = Sci 2009; 00: 000=96000)

(Received July 08, 2009/Revised August 03, = 2009/Accepted=20 August 03, 2009)

DIGITAL OBJECT IDENTIFIER (DOI)

10.1111/j.1349-7006.2009.01308.x About=20 DOI

=20

=20

Article Text

Our understanding of biology and genetics of = gliomas has=20 greatly improved during the past two decades, but unfortunately, = this has=20 not yet led to an effective therapy. Treatment with cytostastic = agents and=20 radiotherapy still dominates first-line adjuvant therapy but = targeted=20 intervention is now increasingly being tested, particularly with=20 inhibitors of neo-angiogenesis and growth factor receptors. At the = same=20 time, large scale sequencing of the glioblastoma genome has led to = the=20 discovery of novel genetic alterations and signaling pathways = which may=20 add additional targets for novel = interventions.⁽^1,2^{)=20
This review summarizes genetic alterations operative in the =
development of=20
astrocytic and oligodendroglial gliomas, with an updated model of=20
signaling pathways and the role of glial precursor cell=20
lineages.}

=20

=20

Definition of Glioma Types

Gliomas are the most common primary brain tumors, = accounting=20 for more than 70% of all primary central nervous system (CNS) = neoplasms.=20 The most frequent and most malignant gliomas are glioblastomas = (World=20 Health Organization [WHO] grade IV).⁽³⁼⁹⁶⁵⁾

Primary and secondary = glioblastoma. The=20 majority of glioblastomas (>90%) develop very rapidly in = elderly=20 patients (mean, 62 years) after a short clinical history, = without=20 clinical or histological evidence of a pre-existing, less = malignant=20 precursor lesion (primary or de novo = glioblastoma).=20 Two-thirds of patients with primary glioblastoma have a clinical = history=20 of <3 months.⁽⁴⁾= =20 Secondary glioblastomas are less frequent (<10%), affect = younger=20 patients (mean age at diagnosis, 45 years), and develop more = slowly=20 through progression from low-grade diffuse astrocytoma (WHO grade = II) or=20 anaplastic astrocytoma (WHO grade III). In a population-based = study, the=20 mean time of progression from low-grade glioma to glioblastoma was = 5.3 years and from anaplastic astrocytoma to glioblastoma=20 1.4 years.⁽⁵⁾= =20 Although histologically largely indistinguishable, primary and = secondary=20 glioblastomas develop through different genetic=20 pathways.⁽^4,6⁾

Low-grade diffuse = astrocytoma (WHO=20 grade II) is a well-differentiated and slow-growing tumor, but = shows a=20 consistent tendency to diffusely infiltrate surrounding brain = structures.=20 Therefore, tumors tend to recur after surgical resection and this = is often=20 associated with progression to more malignant histologic types, = that is=20 anaplastic astrocytoma (WHO grade III) and eventually secondary=20 glioblastoma (WHO grade IV).⁽^3,5⁾

Oligodendroglioma (WHO = grade II) is a=20 well-differentiated, slowly growing, diffusely infiltrating tumor = in=20 adults, typically located in the cerebral hemispheres and composed = predominantly of cells morphologically resembling=20 oligodendroglia.⁽³⁾= =20 Anaplastic oligodendroglioma (WHO grade III) is an = oligodendroglioma with=20 focal or diffuse histological features of malignancy and a less = favorable=20 prognosis.⁽³⁾=

Oligoastrocytoma (WHO grade = II) is=20 composed of a conspicuous mixture of two distinct neoplastic cell = types=20 morphologically resembling the tumor cells in oligodendrogliomas = and=20 low-grade astrocytomas.⁽³⁾= =20 Anaplastic oligoastrocytoma (WHO grade III) is an oligoastrocytoma = with a=20 histological feature of malignancy. With respect to gene status = and=20 survival, oligoastrocytomas are intermediate between low-grade = diffuse=20 astrocytomas and oligodendrogliomas.⁽^3,7⁾

=20

=20

EGFR/RAS/NF1/PTEN/PI3K Pathway

Growth receptors (e.g. EGFR, PDGFRA) become = activated=20 through the binding of their respective ligands (e.g. EGF, = TGF-α, PDGF) to=20 their extracellular domain, which results in recruitment of=20 phosphatidylinositol 3-kinase (PI3K) to the cell membrane. The = PI3K=20 complex is composed of a catalytically active protein p110α = (encoded by=20 PIK3CA) and a regulatory protein p85α = (encoded by=20 PIK3R1). PI3K = phosphorylates=20 phosphatidynositol-4,5-bisphosphate (PIP₂) = to the=20 respective 3-phosphate (PIP₃) which = activates=20 downstream effector molecules such as AKT (protein kinase B) and = mTOR, the=20 mammalian target of rapamycin. This results in cell proliferation = and=20 increased cell survival by blocking apoptosis. PTEN inhibits the=20 PIP₃ signal,⁽⁸⁾= =20 thereby inhibiting cell proliferation (Fig. 1a).=20 The NF1 tumor suppressor gene encodes = neurofibromin=20 that functions primarily as a RAS negative regulator, and also = plays a=20 role in adenylate cyclase and AKT=96mTOR mediated pathways (Fig. 1a).⁽⁹⁾=

=20

= Fig. 1. = Major=20 signaling pathways involved in the pathogenesis of = glioblastomas.=20 pGBM, primary glioblastoma; sGBM, secondary glioblastoma; = α-KG,=20 α-ketoglutarate; FH, fumarase; SDH, succinate = dehydrogenase; HIF-1,=20 hypoxia-inducible factor-1; VHL, Von Hippel=96Lindau. =

[Normal View=20 ]

EGFR amplification occurs = in=20 approximately 40% of primary glioblastomas, but is very rare in = secondary=20 glioblastomas.⁽^4,10,11⁾⁼=20 All primary glioblastomas with EGFR = amplification=20 showed EGFR overexpression, while 70=9690% = of those=20 with EGFR overexpression had gene=20 amplification.⁽^12,13^{)<=
/SUP>EGFR amplification is often associated with =
deletion=20
mutants, EGFRvIII (deletion of exons 2=967) =
being the=20
most frequent type. EGFRvIII shows =
constitutive=20
activation of the receptor, and failure to attenuate signaling by =
receptor=20
down-regulation, and exerts mitogenic =
effects.⁽¹⁴⁾}

The PTEN gene is mutated in = 15=9640% of=20 primary glioblastomas, but rare in secondary=20 glioblastomas.⁽^4,12,15⁾⁼PIK3CA mutations and amplification are rare (5% = and 13%) in=20 both primary and secondary = glioblastomas.⁽¹⁶^{)=20
About two-thirds (63%) of primary glioblastomas and one-third of =
secondary=20
glioblastomas showed alterations in at least one of EGFR,=20
PTEN, or PIK3CA=20
genes.⁽¹⁶^{)=20
The Cancer Genome Atlas (TCGA) pilot project identified additional =
genetic=20
alterations in (mostly primary) glioblastomas, that is NF1 mutations/homozygous deletions (18%) and =
PIK3R1 mutations (10%), with overall frequency of =

alterations in the EGFR/RAS/NF1/PTEN/PI3K =
pathway in=20
88% of glioblastomas.⁽¹⁾=
=20
Increased mRNA expression of PDGFR has been observed in astrocytic =
tumors=20
of all grades, but gene amplification was only detected in a small =
subset=20
of glioblastomas.⁽¹⁷⁾}}

About 50% of both oligodendrogliomas and = anaplastic=20 oligodendrogliomas show EGFR overexpression at the mRNA and = protein=20 levels.⁽¹⁸^{)=20
PDGF A and B, as well as their corresponding receptors =
(PDGFR-α and=20
PDGFR-β) are co-expressed in most=20
oligodendrogliomas.⁽¹⁹^{)=20
In contrast, EGFR amplification, PTEN mutations, and PIK3CA =
mutations=20
may be observed in only a small fraction of anaplastic=20
oligodendrogliomas.⁽³⁾=}}

=20

=20

TP53/MDM2/MDM4/p14^ARF=20 Pathway

The TP53 gene encodes a = protein that=20 plays a role in several cellular processes, including the cell = cycle,=20 response of cells to DNA damage, cell death, and cell=20 differentiation.⁽²⁰^{)=20
Following DNA damage, TP53 is activated and induces transcription =
of genes=20
such as =
p21^Waf1/Cip1.⁽^21,22^{)<=
/SUP>MDM2=20
is induced by wild-type TP53,⁽^23,24^{)<=
/SUP>=20
which binds to mutant and wild-type TP53, thereby inhibiting the =
ability=20
of wild-type TP53 to activate =
transcription.⁽^25,26^{)<=
/SUP>=20
The p14^ARF binds to MDM2 and inhibits=20
MDM2-mediated TP53 degradation and transactivational=20
silencing.⁽^22,27^{)<=
/SUP>=20
MDM4 (also called MDMX) also regulates TP53=20
activity.⁽²⁸^{)=20
p14^ARF is negatively regulated by TP53 (Fig. 1a).⁽²²^{)=20
Thus, loss of normal TP53 function may result from alterations in =
TP53,=20
MDM2, MDM4, or p14^ARF.}}}}}}}

TP53 mutations are = significantly more=20 frequent in secondary glioblastomas than in primary glioblastomas=20 (65%vs 28%).⁽^4,5^{)=20
Of the TP53 mutations in secondary =
glioblastomas, 57%=20
were in hot-spot codons 248 and 273, while in primary =
glioblastomas,=20
mutations were more evenly distributed. G:C→A:T mutations at =
CpG sites=20
were significantly more frequent in secondary than primary =
glioblastomas,=20
suggesting that the acquisition of TP53 =
mutations in=20
these glioblastoma subtypes may occur through different=20
mechanisms.⁽⁴⁾=
=20
Promoter methylation of p14^ARF is frequent in secondary =
glioblastomas,=20
and the analysis of multiple biopsies from the same patients =
revealed=20
p14^ARF=20
methylation already in one-third of low-grade=20
astrocytomas.⁽²⁹^{)=20
Loss of p14^ARF expression due to =
homozygous=20
deletion or promoter methylation is frequent in primary =
glioblastomas=20
(50%).⁽²⁹^{)MDM2 amplification is not frequent=20
(<15%);⁽²⁹^{)=20
it occurs exclusively in primary glioblastomas that lack a TP53 mutation.⁽^30,31^)<=
/SUP>}}}}

At least one alteration in the TP53/MDM2/p14^ARF pathway = (TP53 mutations, p14^ARF homozygous deletion, p14^ARF=20 promoter methylation, MDM2 amplification) = was=20 observed in approximately 50% of primary glioblastomas, and in = >70% of=20 secondary glioblastomas.⁽³²^{)=20
There was a significant inverse correlation between MDM2 amplification and TP53 mutation=20
and between p14^ARF alterations and TP53=20
mutations.⁽³²^{)=20
The recent TCGA pilot project (mostly primary glioblastomas) =
showed that=20
the overall frequency of genetic alterations in the=20
TP53/MDM2/MDM4/p14^ARF pathway in =
glioblastomas was=20
87%, through TP53 mutations or homozygous =
deletion=20
(35%), MDM2 amplification (14%), MDM4 amplification (7%), or p14^ARF=20
homozygous deletion or mutation (49%).⁽¹⁾=}}

Approximately 20% of oligodendrogliomas showed = p14^ARF=20 promoter methylation,⁽³³^{)=20
and p14^ARF homozygous deletion or promoter=20
methylation were observed in 40% of anaplastic=20
oligodendrogliomas.⁽³³^{)=20
The overall frequencies of alterations in the=20
TP53/MDM2/p14^ARF pathway were 21% in=20
oligodendrogliomas and 50% in anaplastic=20
oligodendrogliomas.⁽³³⁾}}

=20

=20

p16^INK4a/CDK4/RB1=20 Pathway

The RB1 protein controls the progression through=20 G₁ into the S-phase of the cell cycle. The = CDK4/cyclin D1 complex phosphorylates the RB1 protein, thereby = inducing=20 release of the E2F transcript factor that activates genes involved = in the=20 G₁→S = transition.⁽²¹^{)=20
p16^INK4a binds to CDK4, inhibits the =
CDK4/cyclin=20
D1 complex, and thus inhibits the =
G₁→S transition=20
(Fig. 1a).⁽²¹^{)=20
Therefore, loss of normal RB1 function may result from altered =
expression=20
of any of the p16^INK4a, CDK4, =
or RB1 genes.}}

Homozygous deletion of the p16^INK4a gene,=20 CDK4 amplification, and loss of RB1 were = largely=20 mutually exclusive; the overall frequency of these alterations was = 50% in=20 primary glioblastomas and approximately 40% in secondary=20 glioblastomas.⁽³⁴^{)=20
The recent TCGA pilot project (mostly primary glioblastomas) =
showed that=20
the overall frequency of genetic alterations in the RB1 signaling =
pathway=20
was 78%, through p16^INK4a homozygous deletion or =
mutations (52%),=20
p15^INK4b=20
homozygous deletion (47%), p18^INK4c homozygous deletion (2%), =
CDK4 amplification (18%), cyclin =
D₂=20
(CCND2) amplification (2%), CDK6=20
amplification (1%), RB1 mutation or =
homozygous=20
deletion (11%).⁽¹⁾=}

Alterations in the p16^INK4a/CDK4/RB1 pathway=20 were rare (4%) in oligodendrogliomas, but frequent (65%) in = anaplastic=20 oligodendrogliomas.⁽³³⁾

=20

=20

IDH1

The IDH1 gene at 2q33 = encodes=20 isocitrate dehydrogenase 1 (IDH1),⁽³⁵^{)=20
which catalyzes the oxidative carboxylation of isocitrate to=20
α-ketoglutarate, resulting in the production of NADPH in the =
citric acid=20
(Krebs) cycle (Fig. 1b).⁽^36,37^{)<=
/SUP>=20
In contrast to other mitochondria IDH forms, IDH1 is present in =
the=20
cytosol. IDH1 mutations decrease the enzyme =

activities in =
vitro.⁽³⁸^{)=20
Heterozygous IDH1 mutations impair the =
enzyme's=20
affinity for its substrate and dominantly inhibit wild-type IDH1 =
activity=20
through the formation of catalytically inactive=20
heterodimers.⁽³⁹^{)=20
Forced expression of mutant IDH1 in cultured cells reduced =
formation of=20
the enzyme product, α-ketoglutarate (α-KG), and =
increased the levels of=20
hypoxia-inducible factor subunit HIF-1α, a transcription =
factor that=20
facilitates tumor growth (Fig. 1b).⁽³⁹⁾}}}}

IDH1 mutations were first = identified in=20 an analysis of 20 661 protein-coding genes in=20 glioblastomas.⁽²⁾= =20 Subsequent studies demonstrated that IDH1 = mutations=20 are very frequent (>80%) in low-grade astrocytomas, anaplastic=20 astrocytomas, secondary glioblastomas, oligodendrogliomas, = anaplastic=20 oligodendrogliomas, oligoastrocytomas, and anaplastic=20 oligoastrocytomas.⁽^38,40,41⁾=20 In contrast, IDH1 mutations appear to be = rare=20 (<5%) or absent in pilocytic astrocytomas and primary=20 glioblastomas,(^38,40,41⁾=20 and absent in ependymomas and other CNS tumors, as well as in = tumors=20 outside of the nervous system (Fig. 2).(^38,40=9642⁾

=20

= Fig. 2. = IDH1 mutations are frequent genetic = alterations in=20 astrocytomas and oligodendrogliomas, very rare in other = tumors of=20 the nervous system, and virtually absent in tumors outside = the=20 central nervous system (CNS). Data on CNS tumors are = combined from=20 Balss et al.,⁽⁴⁰^{)=20
Watanabe et al.,⁽⁴¹^{)=20
and Yan et al.;⁽³⁸^{)=20
data on other tumors including cancers of the bladder, =
breast,=20
stomach, colorectal, lung, thyroid, ovary, and prostate; =
lymphoma=20
and leukemia; and melanoma, are combined from Yan et al.⁽³⁸^{)=20
and Bleeker et al.⁽⁴²⁾⁼²⁰}}}}

[Normal View=20 ]

The majority (>60%) of low-grade astrocytomas = have TP53 mutations plus IDH1=20 mutations,(⁴¹^{)=20
while the majority of oligodendrogliomas typically show IDH1 mutations plus 1p/19q loss (>60%). IDH1 mutations are therefore the first and only =
genetic=20
alterations that are frequent among diffuse astrocytomas,=20
oligodendrogliomas, and oligoastrocytomas, suggesting that these =
gliomas=20
may share the same cell of origin (Fig. 3).=20
The histological criteria for the diagnosis of low-grade =
astrocytomas,=20
oligodendrogliomas, and, in particular, oligoastrocytomas are =
highly=20
subjective. Since the majority of these low-grade gliomas are now=20
genetically characterized by the presence of an IDH1=20
mutation plus TP53 mutation or of an IDH1 mutation plus 1p/19q loss, the molecular=20
classification of low-grade diffuse gliomas may eventually replace =
the=20
histological classification.}

=20

= Fig. 3. = Proposed=20 genetic pathways to astrocytic and oligodendroglial = gliomas.=20

[Normal View=20 ]

=20

=20

Loss of Heterozygosity (LOH)

The most frequent genetic alteration in primary=20 glioblastomas (up to 80%) is LOH on chromosome 10, often with loss = of an=20 entire allele (10p and 10q).⁽^4,43=9646⁾=20 There are at least three commonly deleted loci, that is 10p14-15, = 10q23-24=20 (PTEN), and 10q25-pter, suggesting the = presence of=20 several tumor suppressor genes that may play roles in their=20 pathogenesis.(^43=9645,47⁾=20 LOH 10q is also frequent in secondary glioblastomas (up to=20 70%),(^4,46^{)=20
although it is usually partial; LOH at 10q25-qter was associated =
with=20
histologically-recognized progression from low-grade or anaplastic =

astrocytoma to a highly anaplastic glioblastoma phenotype. LOH 10p =
is very=20
rare in secondary glioblastomas.⁽⁴⁶^{)=20
LOH 10q has been reported in 35=9660% of anaplastic=20
astrocytomas.⁽^45,48^{)<=
/SUP>=20
LOH 10q is infrequent occurring in only approximately 10% of =
anaplastic=20
oligodendrogliomas.⁽⁴⁹⁾}}}

LOH 13q has been detected in 12% of primary and in = 38% of=20 secondary glioblastomas and typically includes the RB1 locus.⁽⁵⁰^{)=20
LOH 6q occurs in approximately one-third of anaplastic=20
astrocytomas.⁽⁵¹^{)=20
The chromosomal region at 9p23-24.1 is frequently deleted in=20
glioblastomas. It encodes the PTPRD gene, a =
receptor=20
protein tyrosine phosphatise with tumor suppressor function that =
is also=20
mutated in a small subset (6%) of glioblastomas and frequently =
(37%)=20
inactivated by promoter methylation.⁽^52,53^)<=
/SUP>}}

LOH 22q is frequent in primary glioblastomas (41%) = and in=20 secondary glioblastomas (82%).⁽⁵⁴^{)=20
Characterization of the 22q deletions in primary glioblastomas =
identified=20
two minimally deleted regions at 22q12.3-13.2 and=20
22q13.31.⁽⁵⁴^{)=20
The small (957 kb) deletion was identified in 22 of 23 =
secondary=20
glioblastomas, a region in which the tissue inhibitor of=20
metalloproteinases-3 (TIMP-3) is =
located.⁽⁵⁴^{)=20
LOH on 22q has been reported at a frequency of 20=9630% in both =
low-grade=20
and anaplastic astrocytomas.⁽³⁾=}}}

=20

=20

Loss of 1p and 19q

Oligodendrogliomas and anaplastic = oligodendrogliomas=20 typically show concurrent deletion of chromosomes 1p and 19q (up = to 80%),=20 and this is associated with increased chemosensitivity to = treatment with=20 PCV (procarbazine, CCNU, and vincristine) and a more favorable = clinical=20 outcome.⁽^55=9658⁾=20 In most cases, the entire 1p/19q arms are=20 involved.(^57,59=9661⁾=20 Jenkins et al.(⁵⁸^{)=20
showed that combined deletion of chromosomes 1p and 19q is due to=20
unbalanced translocation between chromosomes 1 and 19 =
[t(1;19)(q10;p10)].=20
The prevalence of this translocation suggests that the combined =
loss of=20
two or more genes on 1p and 19q are required for the development =
of the=20
majority of oligodendrogliomas.⁽⁵⁸^{)=20
Molecular cytogenetic deletion mapping studies have suggested that =
the=20
minimal regions of deletion and, by implication, the putative =
candidate=20
genes reside within 1p36 and 19q13.3.⁽^57,61,62⁾=20
Isolated deletions of 19q are also common in astrocytic and=20
oligodendroglial tumors.(}^61,63,64⁾=20
Isolated deletions of 1p are rare in gliomas and are associated =
with a=20
poorer prognosis.(}^57,65^)<=
/SUP>

LOH 1p is rare in both primary (12%) and secondary = glioblastomas (15%),⁽⁵⁰^{)=20
but is associated with longer patient =
survival.⁽⁶⁶^{)=20
LOH 19q (common deletion at 19q13.3) has been found to be frequent =
in=20
secondary glioblastomas (54%) but rare in primary glioblastomas=20
(6%).⁽⁵⁰⁾}}

=20

=20

cDNA and Protein Expression = Profiles

Although usually histologically indistinguishable, = primary=20 and secondary glioblastomas show significantly different mRNA and = protein=20 expression profiles, reflecting their significant difference in = genetic=20 alterations.⁽⁶⁾= =20 Genes typically expressed in primary glioblastomas include=20 Fas,⁽⁶⁷^{)=20
VEGF,⁽⁶⁸^{)=20
and VEGF fms-related tyrosine kinase 1, which are likely a =
reflection of=20
the presence of large ischemic necroses in primary=20
glioblastomas.⁽⁶⁷^{)=20
Insulin-like growth factor binding protein-2 (IGFBP-2), a =
modulator of the=20
action of insulin-like growth factors, is also typically expressed =
at a=20
high level in primary glioblastoma.⁽⁶⁹^{)=20
Primary glioblastomas also preferentially express genes typical of =
a=20
stromal response, suggesting the importance of extracellular=20
signaling.⁽⁷⁰^{)AEBP1,=20
up-regulated >4-fold in the majority of primary glioblastomas =
compared=20
to secondary glioblastomas, is a transcriptional repressor, and =
has been=20
shown to interact with PTEN and inhibit its=20
function.⁽⁷¹^{)=20
Proteins expressed uniquely in primary glioblastomas include =
tenascin-X=20
precursor, enolase 1, centrosome-associated protein 350, and EGFR. =
A=20
significant variation in cDNA expression profiles was observed =
among=20
different primary glioblastomas, probably reflecting their genetic =

heterogeneity.⁽⁶⁹⁾}}}}}}

ASCL1 is overexpressed in 86% of low-grade = astrocytomas and=20 88% of secondary glioblastomas, which is accompanied by inhibition = of=20 Notch signaling.⁽⁷²^{)=20
Other proteins typically expressed in secondary glioblastomas =
include=20
ERCC6, DUOX2, HNRPA3, WNT-11 protein precursor, cadherin-related =
tumor=20
suppressor homolog precursor, and =
ADAMTS-19.⁽⁷³^{)=20
Quantification of the cytokines in the supernatant of 30=20
tissue-correspondent glioma cultures revealed a high expression =
level of=20
PDGF-AB in secondary glioblastomas.⁽⁶⁸^{)=20
The extent of variation of cDNA expression profiles among =
low-grade=20
astrocytomas was much less than that among glioblastomas, =
suggesting that=20
low-grade astrocytomas are genetically more=20
homogeneous.⁽^69,74^)<=
/SUP>}}}

=20

=20

Response to Chemotherapy

In most centers, first-line adjuvant therapy for=20 glioblastomas consists of treatment with the alkylating agent = temozolomide=20 (TMZ) and concurrent radiotherapy.⁽^75,76^{)<=
/SUP>O⁶-Methylguanine-DNA =
methyltransferase (MGMT)=20
is a repair protein that specifically removes promutagenic alkyl =
groups=20
from O⁶-methylguanine,⁽⁷⁷^{)=20
a promutagenic DNA adduct that causes G:C→A:T mutations =
during DNA=20
replication.⁽^78,79^{)<=
/SUP>=20
Loss or reduction of MGMT activity due to promoter methylation is =
frequent=20
(40=9657%) in glioblastomas,⁽^80=9682⁾=20
and MGMT methylation is associated with =
response to=20
chemotherapy and longer survival among glioblastoma patients =
treated with=20
TMZ.(}^80,83,84⁾}}

=20

=20

Chemotherapy-induced Genetic = Alterations

Some genetic alterations appear to be associated = with=20 chemotherapy. Mutations and loss of expression of the MSH6 mismatch repair gene have been observed in a = small=20 subset of glioblastomas treated with TMZ, but not in untreated=20 glioblastomas.(⁸⁵^{)MSH6 mutations were significantly associated with =
a=20
hypermutator glioblastoma phenotype.⁽^75,85^{)<=
/SUP>=20
In the recent TCGA study, seven hypermutated tumors were observed; =
all=20
were from patients previously treated with TMZ or CCNU. Six of the =
seven=20
hypermutated glioblastomas harbored mutations in at least one of =
the=20
mismatch repair genes MLH1, MSH2, MSH6, or PMS2, as compared with only one sample among 84=20
non-hypermutated cases.⁽¹⁾=}}

=20

=20

Brain Tumor Stem Cells

Most genetic analyses in brain tumors were carried = out on=20 the bulk tumor mass. The cancer stem cell hypothesis postulates = that only=20 a small fraction of tumor cells have the capacity for self-renewal = and=20 tumor initiation. These brain tumor stem cells differ from the = majority of=20 tumor cells in gene expression⁽⁸⁶^{)=20
and biological behavior, including proliferation, spread, and =
sensitivity=20
to chemo- and radiotherapy.⁽^87,88^)<=
/SUP>}

CD133 (prominin) has been used as a putative stem = cell=20 marker in normal and malignant brain tissues. Brain tumor cells = expressing=20 CD133 appear to possess a capacity for proliferation, = self-renewal, and=20 differentiation.⁽⁸⁹^{)=20
Only the CD133+ brain tumor fraction contained cells that were =
capable of=20
tumor initiation in NOD-SCID mouse =
brains,⁽⁹⁰^{)=20
although this observation was not confirmed in another=20
study.⁽⁹¹^{)=20
CD133+ expression in glioblastoma cells was associated with their=20
resistance to radiotherapy and =
chemotherapy.⁽^87,88^{)<=
/SUP>=20
CD133+ and CD133− glioblastoma-derived cancer stem cells =
showed=20
differential growth characteristics and molecular=20
profiles.⁽⁸⁶^{)=20
One study showed that the relative content of CD133+ cells was=20
significantly higher in primary than in secondary glioblastomas, =
and that=20
CD133+ expression was associated with neurosphere formation only =
in=20
primary glioblastomas,⁽⁸⁶^{)=20
reinforcing the view that the origin and cancer stem cells in =
primary and=20
secondary glioblastomas may be different. Alternatively, CD133 may =
not be=20
the only brain stem cell marker. Griguer et al.⁽⁹²^{)=20
reported evidence that CD133 expression may be reversibly =
up-regulated by=20
hypoxia or by mitochondrial dysfunction through pharmacological =
inhibition=20
of the electron transport chain.}}}}}}}

=20

=20

Outlook

During the past decade, much progress has been = made in our=20 understanding of the origin, biology, and genetics of gliomas.=20 Regrettably, this has only led to moderate improvements in = clinical=20 outcome. Therapeutic targeting of pathways operative in malignant=20 astrocytomas, for example the EGFR/RAS/NF1/PTEN/PI3K pathway, has = had=20 limited success so far, and in most centers, the first-line = therapy for=20 glioblastoma is still a cytotoxic agent (TMZ) and radiotherapy. = However,=20 we anticipate that the genome-wide sequencing of gliomas will lead = to the=20 identification of additional signaling pathways and novel targets = for=20 therapy. Eventually, extensive genetic profiling of gliomas will = become a=20 laboratory routine, constituting for an important further step = towards=20 personalized treatment.

=20

=20

=20

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