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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_0000_01CAD4A7.54BB7A00 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('
3D"provided
');=0A= document.writeln('
    ');=0A= =0A= var i;=0A= //Change start for bug#2288=0A= if(typeof(maxBranding) =3D=3D "undefined" | maxBranding =3D=3D "" | = maxBranding =3D=3D 0){=0A= maxBranding =3D 3; // DEFAULT Value for No of Cobranding dispaly=0A= }=0A= var size =3D customerCoBranding.length;=0A= if(maxBranding !=3D -1 & size-1 > maxBranding){=0A= size =3D maxBranding;=0A= }=0A= //Change End for bug#2288=0A= =0A= // Add the co-branding images o the page.=0A= //for(i=3D0; i < customerCoBranding.length; i++ )=0A= for(i=3D0; i < size; i++ )=0A= {=0A= // ignore the terminating array element.=0A= if(customerCoBranding[i].image =3D=3D "none")=0A= {=0A= continue;=0A= }=0A= =0A= this.image =3D customerCoBranding[i].image;=0A= this.imageAltText =3D customerCoBranding[i].imageAltText;=0A= this.href =3D customerCoBranding[i].href;=0A= =0A= if (image =3D=3D "")=0A= {=0A= if(href =3D=3D "")=0A= {=0A= document.writeln('
  • ' + imageAltText + '
    • ');=0A= }=0A= else=0A= {=0A= document.writeln('
  • ' + imageAltText + = '
    • ');=0A= }=0A= }=0A= else=0A= {=0A= if(href =3D=3D "")=0A= {=0A= document.writeln('
  • 3D"'
    • ');=0A= }=0A= else=0A= {=0A= document.writeln('
  • 3D"'
    • ');=0A= }=0A= }=0A= }=0A= =0A= // If there are more co-branding images add in a link to them.=0A= // Changes made for Bug#2288=0A= //if(moreCustomerCoBranding =3D=3D "true")=0A= if(maxBranding !=3D -1 & customerCoBranding.length-1 > maxBranding)=0A= {=0A= document.writeln('
  • More Providers
    • ');=0A= }=0A= =0A= document.writeln('
');=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("
    ");=0A= if(typeof(environment)!=3D'undefined' && environment!=3D'PRODUCTION')=0A= {=0A= if(userName !=3D "")=0A= {=0A= //write out the user specific data for the menu=0A= if (userdata !=3D null){=0A= if (userName.length > 33){=0A= var writedata =3D "
  • " + = userName.substring(0,33) + "..., access your
    • ";=0A= } else {=0A= var writedata =3D "
  • " + userName + = ", access your
    • ";=0A= }=0A= =0A= for (var x=3D0; xMy 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("
    ");=0A= if (userdata !=3D null){=0A= var writedata =3D "";=0A= for (var x=3D0; xLog 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('
  • Login
    • ');=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 "
  • Log Out
    • ";=0A= }else=0A= {=0A= return "
  • Log = In
    • ";=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 "
  • Athens Log In
    • ";=0A= }=0A= =0A= } ------=_NextPart_000_0000_01CAD4A7.54BB7A00 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 "
  • more...
  • " + liels[j].innerHTML + "
    • ";=0A= } else {=0A= tmpinnerhtml +=3D "
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    3D"Jump

    STEM=20 CELLS
    ISSN: 1549-4918 (Online)
    ISSN: = 1066-5099=20 (Print)

    Volume 28, Issue 1, 2010.
    Pages: 5-16
    • Top of page
    • Abstract
    • INTRODUCTION
    • MATERIALS AND METHODS
    • RESULTS
    • DISCUSSION
    • SUMMARY
    • Acknowledgements
    • References
    List of Images
    Abstract=20 Only

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http://www3.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=3D 0) { appname =3D "ns"; }=0A= else if (navigator.appName.indexOf("Explorer") >=3D 0) { appname =3D = "ie"; }=0A= if (navigator.userAgent.indexOf("Mozilla/2") >=3D 0) {=0A= version =3D 2;=0A= } else if (navigator.userAgent.indexOf("Mozilla/3") >=3D 0) {=0A= version =3D 3;=0A= } else if (navigator.userAgent.indexOf("Mozilla/4") >=3D 0) {=0A= version =3D 4;=0A= } else if (navigator.userAgent.indexOf("Mozilla/5") >=3D 0) {=0A= version =3D 5;=0A= }=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= // Get the cookies named "REF_LNX and SRC_LNX"... deprecated for now. = Always set.=0A= // Instead set the hypertext for linking.=0A= //var anchorString =3D '3D"extended';=0A= var anchorString =3D 'Links';=0A= var targString =3D ' target=3D_top';=0A= var anchor2 =3D 'links';=0A= =0A= //NEW for special content linking. Anchor string must be taken from SGML.=0A= var contAnchor =3D '';=0A= =0A= // Linking functions=0A= =0A= =0A= //Get the EarlyView flag of the citing article which is needed to ensure = that the link is suitable for ISI WoS.=0A= // The CONTROL details are passed in, in the form -----. only grab the = 2nd of 5 characters which is the EarlyView flag=0A= function setEarlyView(object)=0A= {=0A= if ((version >=3D 3) && (object.substring(1,2) !=3D "Y"))=0A= {=0A= artEarlyView =3D "EVIEW=3DN"; =0A= }=0A= }=0A= =0A= //Get the DOI of the citing article which is needed for the ISI links.=0A= function setDOI(object){=0A= if (version >=3D 3)=0A= {=0A= artDOI =3D object;=0A= }=0A= }=0A= =0A= function setISSN(object)=0A= {=0A= }=0A= =0A= // Print out the reference link. =0A= function print_link(object)=0A= {=0A= if (version >=3D 3)=0A= {=0A= // remove over-escaped ampersands (ref bugzilla 5515)=0A= var overEscapedAmpersandsRemoved =3D object.replace(new = RegExp('&','g'), '&');=0A= =0A= // remove "BIB" prefix from the value of BIBID (for ISI linking)=0A= // note, this could be performed in LNX. Need to reassess where this = should be done.=0A= var re =3D new RegExp ('BIBID=3DBIB', 'i') ;=0A= var paramstring =3D new String();=0A= paramstring =3D overEscapedAmpersandsRemoved.replace(re, 'BIBID=3D');=0A= =0A= var unpCookie =3D '/cgi-bin/reflink?object=3D' + escape(paramstring = + "&" + artDOI + "&" + artEarlyView + "&" + artEnableISI + "&" + = artTurnOnCas + "&" + artEnablePubMed);=0A= if (netscape4 =3D=3D false)=0A= {=0A= document.write ('' + anchorString + '  ');=0A= }=0A= else // protect against inline style application hanging NN4=0A= {=0A= document.write ('' + = anchorString + '  '); =0A= }=0A= }=0A= else=0A= {=0A= document.writeln();=0A= } =0A= }=0A= =0A= //Print out source links (in the citing article)=0A= function print_SRClink(object){=0A= if (version >=3D 3)=0A= {=0A= var unpCookie =3D '/cgi-bin/srclink?object=3D' + escape(object);=0A= var url =3D unpCookie;=0A= document.write ('
  • Find = other ');=0A= document.write (anchor2.link(url));=0A= document.write (' for this article.
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    3D"About

    Online ISSN: 1549-4918    Print ISSN: = 1066-5099
    STEM CELLS
    Volume 28, Issue 1, 2010. Pages: 5-16

    Copyright © 2009 AlphaMed Press

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    Copyright © 2010 AlphaMed = Press
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    '); } =0A= function otherlinks(hwnd, iname, size,id) {hwnd.document.write('View Normal Image '); else = hwnd.document.write('(1) target=3Dmain >View Magnified Image '); if = (id !=3D 'null') hwnd.document.write('Interactive View '); = hwnd.document.write('View List of Images '); hwnd.document.write('

    '); } =0A= function caption_fig001(hwnd) { hwnd.document.write('Figure 1. GBM neurospheres are sensitive to NOTCH pathway blockade in vitro. (A): The GBM neurosphere line HSR-GBM1 grows as large, = multicellular spheres. The majority (70%) of neurosphere cells express CD133. (B): Upon = GSI-18 treatment, protein levels of HES1, a NOTCH pathway target gene, = were reduced after 48 hours of GSI-18 treatment of HSR-GBM1 = neurospheres. (C, D): Upon GSI-18 treatment, mRNA = expression of NOTCH target gene HES5 and protein levels of cleaved = active NOTCH1 intracellular domain are also reduced. (E): = GSI-18 reduced total cell mass in a dose-dependent fashion in all three = HSR-GBM1 sublines (*, p < .05). (F): There is no = significant change in normal human astrocyte growth when treated with = GSI-18. (G): Growth of a second GBM neurosphere line, HSR-GBM2, = was also inhibited by GSI-18. (H): HES1 mRNA levels fell into a = similar range in primary GBM and tumor-derived neurospheres grown in = culture. (I): A second -secretase inhibitor, MRK-003, could also suppress GBM = neurosphere growth. (J): MRK003 showed potency similar to = GSI-18 in terms of its ability to lower HES1 mRNA levels in tumor = neurospheres. (K, L): Low-passage neurospheres GBM-SC, GBM-KK, = and GBM-DM express the NOTCH target gene HEY1 and had their growth = inhibited by NOTCH blockade. Abbreviations: DMSO, dimethyl sulfoxide; = GAPDH, glyceraldehyde-3-phosphate dehydrogenase; GBM, glioblastoma; = GSI-18, -secretase = inhibitor 18.
    '); } =0A= function gcaption_fig001(hwnd) { hwnd.document.write('AlphaMed Press

    '); } =0A= function display_fig001(size) { var i =3D new Array(); i[0] =3D = 'nfig001'; i[1] =3D 'mfig001'; var hwnd =3D newfigwin(); dochead(hwnd, = ''); copyright(hwnd); otherlinks(hwnd, 'fig001', size,'null'); = imagelink(hwnd, i[size]); gcaption_fig001(hwnd); caption_fig001(hwnd); = doctail(hwnd); hwnd.document.close(); hwnd.focus(); } =0A= function caption_fig002(hwnd) { hwnd.document.write('Figure 2. Activation of NOTCH2 in glioblastoma (GBM) neurospheres increases tumor = growth in vitro. (A): When the active form of NOTCH2 = receptor (NICD2) was stably transfected into HSR-GBM1a cells, receptor = mRNA levels were elevated. (B): NICD2 protein expression was = also increased in NICD2 stably transfected HSR-GBM1 neurospheres = detected by Western blot. GAPDH was used as protein loading control. (C): The NOTCH target gene HES1 was also induced in = NICD2-transfected GBM neurospheres. (D): Cell growth was = significantly increased in NICD2-transfected cells. (E): = Transient expression of NICD2 in HSR-GBM1 neurospheres using adenovirus = vectors also increased cell growth over 2 days. Abbreviations: Ad, = Adenovirus; Beta Gal, -galactosidase; GAPDH, glyceraldehyde-3-phosphate = dehydrogenase; NICD2, NOTCH2 intracellular domain.
    '); } =0A= function gcaption_fig002(hwnd) { hwnd.document.write('AlphaMed Press

    '); } =0A= function display_fig002(size) { var i =3D new Array(); i[0] =3D = 'nfig002'; i[1] =3D 'mfig002'; var hwnd =3D newfigwin(); dochead(hwnd, = ''); copyright(hwnd); otherlinks(hwnd, 'fig002', size,'null'); = imagelink(hwnd, i[size]); gcaption_fig002(hwnd); caption_fig002(hwnd); = doctail(hwnd); hwnd.document.close(); hwnd.focus(); } =0A= function caption_fig003(hwnd) { hwnd.document.write('Figure 3. NOTCH blockade depletes CD133-positive cells and inhibits neurosphere = clonogenicity. (A): CD133 mRNA levels were severely decreased = following GSI-18 treatment of HSR-GBM1 cells. (B): The = percentage of CD133-positive cells in these cultures was also lowered by = NOTCH blockade. (C-E): mRNA levels of the stem cell markers = NESTIN, BMI1, and OLIG2 were also reduced in a dose-dependent fashion = following GSI-18 treatment of HSR-GBM1 neurospheres. (F, G): = Growing HSR-GBM1 cultures treated for 5 days with -secretase inhibitor can = form some neurospheres (right top panel), but when equal numbers of = DMSO- and MRK003-treated cells are seeded singly in methylcellulose, the = latter cannot efficiently form colonies. Abbreviations: DMSO, dimethyl = sulfoxide; GAPDH, glyceraldehyde-3-phosphate dehydrogenase; GBM, = glioblastoma; GSI-18, -secretase inhibitor 18.
    '); = } =0A= function gcaption_fig003(hwnd) { hwnd.document.write('AlphaMed Press

    '); } =0A= function display_fig003(size) { var i =3D new Array(); i[0] =3D = 'nfig003'; i[1] =3D 'mfig003'; var hwnd =3D newfigwin(); dochead(hwnd, = ''); copyright(hwnd); otherlinks(hwnd, 'fig003', size,'null'); = imagelink(hwnd, i[size]); gcaption_fig003(hwnd); caption_fig003(hwnd); = doctail(hwnd); hwnd.document.close(); hwnd.focus(); } =0A= function caption_fig004(hwnd) { hwnd.document.write('Figure 4. NOTCH signaling blockade depletes cells required for glioblastoma (GBM) = propagation in vivo. (A-C): Large xenografts formed at all = eight sites (asterisk) injected with cells in which NOTCH signaling was = active, including nontreated (n =3D 4, A) and vehicle = (DMSO)-treated (n =3D 4) cultures. Mice injected with equal = number of viable cells previously treated with 2 M GSI-18 = (n =3D 6) were unable to efficiently form tumors, with small = lesions developing at only three of six injection sites. (D): = Tumor volume was dramatically reduced in mice engrafted with 2 M = GSI-18-pretreated cells compared with mice engrafted with mock or = DMSO-pretreated cells. (E-G): The small subcutaneous xenografts = that formed from cells pretreated with GSI-18 appeared similar to = control tumors, and expressed abundant amounts of the NOTCH pathway = target HES1 on immunohistochemical analysis. (H, I): = Intracranial xenografts of HSR-GBM1 cells pretreated with DMSO or GSI-18 = often invaded into the subventricular space (H), and = occasionally formed necrotic foci with pseudopalisading (I). = (Original magnifications both =D7100.) Abbreviations: DMSO, dimethyl = sulfoxide; GSI-18, -secretase inhibitor 18.
    '); = } =0A= function gcaption_fig004(hwnd) { hwnd.document.write('AlphaMed Press

    '); } =0A= function display_fig004(size) { var i =3D new Array(); i[0] =3D = 'nfig004'; i[1] =3D 'mfig004'; var hwnd =3D newfigwin(); dochead(hwnd, = ''); copyright(hwnd); otherlinks(hwnd, 'fig004', size,'null'); = imagelink(hwnd, i[size]); gcaption_fig004(hwnd); caption_fig004(hwnd); = doctail(hwnd); hwnd.document.close(); hwnd.focus(); } =0A= function caption_fig005(hwnd) { hwnd.document.write('Figure 5. Local GSI treatment prolongs survival of mice bearing intracranial GBM = xenografts. (A): Wild-type or NICD2 stably transfected GBM = neurospheres were implanted into mouse brain, and GSI-18- or DMSO-soaked = beads were injected into the tumor beds 2 weeks later. Tumor growth was = monitored by MRI in selected animals, and large xenografts formed in = DMSO- but not GSI-18-treated mice. As expected, constitutive NICD2 = expression rendered xenografts insensitive to GSI-18. (B): = GSI-18-treated mice have significantly prolonged survival compared with = DMSO-treated mice, whereas expression of activated NOTCH2 promoted more = rapid intracranial tumor growth. (C-E): Microscopic examination = of a GSI-18-treated mouse brain stained with H&E reveals a round = polymer bead (arrow) with a pale linear injection tract leading upward = (arrowhead), but no tumor mass on this or additional step sections = (original magnification =D7100). Immunocytochemical stains using an = antibody specific for human nuclei confirms the absence of viable tumor = cells (D, E). (Original magnification =D7100.) Abbreviations: = DAPI, 4,6-diamidino-2-phenylindole; DMSO, dimethyl sulfoxide; = GAPDH, glyceraldehyde-3-phosphate dehydrogenase; GBM, glioblastoma; = GSI-18, -secretase = inhibitor 18; Hu, human nuclei; MRI, magnetic resonance imaging; NICD2, = NOTCH2 intracellular domain.
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    3D""=20

    STEM CELLS

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    Volume 28, Issue 1, Pages 5-16

    Published Online: 10 Nov 2009

    Copyright =A9 2010 AlphaMed Press

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    3D""=20
    3D""=20
     Cancer Stem Cells

    *Correspondence to Xing Fan, = Assistant=20 Professor of Neurosurgery and Cell & Developmental Biology, = University=20 of Michigan Medical School, Department of Neurosurgery, 109 Zina = Pitcher=20 Place, 5,018 BSRB, Ann Arbor, Michigan 48109-2200, = USA

    Author contributions: X.F.: conception and = design,=20 financial support, administrative support, provision of study = material,=20 collection and/or assembly of data, data analysis and = interpretation,=20 manuscript writing, final approval of manuscript; L.K., T.S.Z., = M.E.S.,=20 C.E.T., N.G., C.K., J.Z.: collection and/or assembly of data; = Y.-M.L.,=20 J.M., G.N., F.D., S.P., A.L.V.: provision of study material; = C.G.E.:=20 conception and design, financial support, administrative support,=20 provision of study material, data analysis and interpretation, = manuscript=20 writing.
    Disclosure of potential conflicts of interest is = found at=20 the end of this article.
    =A7First published online = in STEM=20 CELLS EXPRESS November 10, = 2009.
    =B6Telephone:=20 734-615-7266; Fax: 734-763-7322

    Funded by:
     Accelerate Brain Cancer Cure Project Award
     American Brain Tumor Association Translational = Grant
     Voices Against Brain Cancer Research Grant; Grant = Number:=20 NIH/NS55089
     Brain Tumor Funders Collaborative

    NOTCH Pathway Blockade Depletes=20 CD133-Positive Glioblastoma Cells and Inhibits Growth of = Tumor=20 Neurospheres and Xenografts =A7
    Xing Fan 1 2=20 *=B6, Leila Khaki=20 3, Thant S. Zhu = 1, Mary=20 E. Soules 1, Caroline E.=20 Talsma 1, Naheed = Gul 3, Cheryl Koh=20 3, Jiangyang Zhang = 4,=20 Yue-Ming Li 8, Jarek=20 Maciaczyk 9, Guido = Nikkhah 9, Francesco DiMeco 7=20 10, Sara Piccirillo = 11,=20 Angelo L. Vescovi 11, = Charles G.=20 Eberhart 3 5 6
    1Department of = Neurosurgery,=20 University of Michigan Medical School, Ann Arbor, Michigan,=20 USA
    2Department of Cell = and=20 Developmental Biology, University of Michigan Medical = School, Ann=20 Arbor, Michigan, USA
    3Department=20 of Pathology, Johns Hopkins University, Baltimore, Maryland, = USA
    4Department of = Radiology,=20 Johns Hopkins University, Baltimore, Maryland, = USA
    5Department of Oncology, Johns = Hopkins=20 University, Baltimore, Maryland, USA
    6Department of Ophthalmology, Johns = Hopkins=20 University, Baltimore, Maryland, USA
    7Department of Neurological Surgery, = Johns=20 Hopkins University, Baltimore, Maryland, USA
    8Pharmacology & Chemistry = Program, Memorial=20 Sloan-Kettering Cancer Center, New York, New York, = USA
    9Department of Stereotactic and = Functional=20 Neurosurgery, University of Freiburg, Freiburg,=20 Germany
    10Department of = Neurosurgery, Istituto Nazionale Neurologico C. Besta, = Milan,=20 Italy
    11Department of=20 Biotechnology and Biosciences, University of Milan Bicocca, = Milan,=20 Italy
    email: Xing Fan (xingf@umich.edu)
    Keywords
    Cancer Stem Cell =95 NOTCH =95 Glioblastoma =95 -Secretase inhibitor

    Abstract
    3DAbstract=20 3D""=20
    Cancer stem cells (CSCs) are thought to be critical for = the=20 engraftment and long-term growth of many tumors, including=20 glioblastoma (GBM). The cells are at least partially spared = by=20 traditional chemotherapies and radiation therapies, and = finding new=20 treatments that can target CSCs may be critical for = improving=20 patient survival. It has been shown that the NOTCH signaling = pathway=20 regulates normal stem cells in the brain, and that GBMs = contain=20 stem-like cells with higher NOTCH activity. We therefore = used=20 low-passage and established GBM-derived neurosphere cultures = to=20 examine the overall requirement for NOTCH activity, and also = examined the effects on tumor cells expressing stem cell = markers.=20 NOTCH blockade by -secretase inhibitors (GSIs) reduced neurosphere = growth and=20 clonogenicity in vitro, whereas expression of an active form = of=20 NOTCH2 increased tumor growth. The putative CSC = markers=20 CD133, NESTIN, BMI1, and OLIG2 were reduced following NOTCH=20 blockade. When equal numbers of viable cells pretreated with = either=20 vehicle (dimethyl sulfoxide) or GSI were injected = subcutaneously=20 into nude mice, the former always formed tumors, whereas the = latter=20 did not. In vivo delivery of GSI by implantation of = drug-impregnated=20 polymer beads also effectively blocked tumor growth, and=20 significantly prolonged survival, albeit in a relatively = small=20 cohort of animals. We found that NOTCH pathway inhibition = appears to=20 deplete stem-like cancer cells through reduced proliferation = and=20 increased apoptosis associated with decreased AKT and STAT3=20 phosphorylation. In summary, we demonstrate that NOTCH = pathway=20 blockade depletes stem-like cells in GBMs, suggesting that = GSIs may=20 be useful as chemotherapeutic reagents to target CSCs in = malignant=20 gliomas. STEM CELLS=20 2010;28:5-16

    Received: 29 May 2009; Accepted: 30 October 2009

    Digital Object = Identifier=20 = (DOI)

    10.1002/stem.254  About=20 DOI

    Article=20 Text

    INTRODUCTION
    3DINTRODUCTION=20 3D""=20

    More than 41,000 people are diagnosed with primary brain tumors = each=20 year in the U.S. [[1]],=20 and glioblastomas (GBMs) are the most common malignant brain = tumors in=20 adults [[2]].=20 Even with advances in surgery, imaging, chemotherapy, and = radiation=20 therapy over the past three decades, more than 70% of GBM patients = still=20 die within 2 years of diagnosis [[1-3]].=20 New strategies to treat this deadly disease are desperately needed = [[1]].=20 In this study, we focus on the potential therapeutic role of NOTCH = pathway=20 inhibition, and the requirement for ongoing NOTCH signaling in = stem-like=20 tumor cells.

    There is emerging evidence that a small population of cancer = stem cells=20 (CSCs) within neoplasms is responsible for long-term tumor = propagation,=20 with initial studies focused on leukemia [[4]].=20 Several groups have subsequently demonstrated that stem-like cells = exist=20 in brain tumors, including GBMs [[4-9]].=20 CSC markers such as CD133 and side population have been used to=20 prospectively isolate a small fraction of cells in human and = rodent brain=20 tumors with a significantly increased potential to generate tumor=20 neurospheres and xenografts [[5][7][8][10][11]].=20 Tumor-derived neurospheres also have a genetic profile that = remains closer=20 to that of the tumor from which they are isolated than traditional = adherent GBM cell lines maintained in high serum [[12]].=20 In addition, GBMs propagated as neurospheres maintain stem-like=20 subpopulations, and more accurately replicate the infiltrating = growth=20 patterns seen in primary tumors [[9][12]].=20 GBM neurosphere cultures thus provide an improved pathological = model, and=20 may also facilitate the discovery of new therapeutic reagents that = can=20 target the stem-like or xenograft-initiating cells required for = long-term=20 tumor growth.

    The Notch locus was first described by Morgan in a = strain of=20 Drosophila with notched wing blades [[13]].=20 The gene was subsequently cloned as a cell surface receptor [[14]],=20 playing a key role in the development of many different cell types = and=20 tissues, including neurons in the central nervous system [[15-18]].=20 NOTCH signaling is initiated when transmembrane ligands on one = cell bind=20 NOTCH receptors on an adjacent cell and cause the -secretase-mediated proteolytic release of the NOTCH=20 intracellular domain (NICD) [[19][20]].=20 NICD then translocates into the nucleus where it interacts with = the=20 transcriptional cofactor CBF1 and activates targets such as the = HES and=20 HEY genes, which modulate neuronal and glial differentiation [[21]].=20 In vertebrates, at least four NOTCH receptors (NOTCH 1-4), five = ligands=20 (JAG1, 2, DLL1, 3, 4), and multiple effector molecules = (HES1-6,=20 HEY1, 2 L) have been identified [[21][22]].=20 NOTCH ligands, receptors, and targets have been found in a wide = range of=20 neoplasms, including lung, breast, cervix, head/neck, renal, and = pancreas=20 carcinoma, neuroblastoma, myeloma, melanoma, choroid plexus tumor, = medulloblastoma, and GBM [[20][22-33]].=20 In many of these tumor types, it has been shown that increased = NOTCH=20 activity promotes tumor growth, whereas NOTCH pathway blockade = inhibits=20 proliferation and/or survival. Inhibition of NOTCH signaling is = therefore=20 a promising therapeutic avenue in a wide range of cancers.

    We and others have demonstrated that the NOTCH signaling = pathway plays=20 an important role in the pathogenesis of medulloblastoma and GBM = [[28][34-36]],=20 and that these malignant brain tumors contain stem-like cancer = cells with=20 higher NOTCH activity [[9][12][32]].=20 It has recently been reported that common chemotherapeutic drugs,=20 including temozolomide, carboplatin, paclitaxel (Taxol), and = etoposide=20 (VP16), as well as traditional radiation therapy, predominantly = targeted=20 the CD133-negative population, and spared or enriched the = CD133-positive=20 population [[37][38]].=20 We have also observed an increase in the stem-like subpopulation = in=20 GBM-derived neurosphere lines following radiation therapy [[39]].=20 Thus, conventional chemotherapies and radiation therapies appear = to=20 effectively remove only better-differentiated cells, while leaving = many=20 GBM CSCs alive. The NOTCH pathway represents a possible target in=20 stem-like glioma cells, as several groups have shown that GBM CSCs = express=20 NOTCH family genes, and that tumor-derived neurospheres have = elevated=20 NOTCH activity [[9][12][32]].=20 It is known that NOTCH regulates normal neural stem cell = self-renewal and=20 differentiation [[40][41]],=20 and that CSCs share many characteristics with their normal = cognates,=20 including the signaling pathways that regulate self-renewal. In = the=20 present study, we used both established and low-passage GBM = cultures as a=20 model to examine the effects of NOTCH pathway blockade on CSCs and = tumor=20 growth.

    MATERIALS AND=20 METHODS
    3D"MATERIALS 3D""=20

    Cell = Culture

    GBM neurosphere cultures were maintained in Neurocult medium = (Stem Cell=20 Technologies, Vancouver, BC, Canada, http://www.stemcell.com/) supplemented with = epidermal=20 growth factor (20 ng/ml) and fibroblast growth factor (10 ng/ml). = Cell=20 pools and stable subclones transfected with NOTCH2 = intracellular=20 domain (NICD2) were generated as previously described [[34][42]].=20 For treatment studies, cells were plated and allowed to grow = overnight in=20 Neurocult medium; Neurocult was then replaced the next morning = with medium=20 containing -secretase inhibitor 18 (GSI-18) dissolved in dimethyl = sulfoxide=20 (DMSO) at the concentrations of 0.4-50 M. RNA and protein extractions and = all cell-based=20 assays were performed 2-5 days after drug application. Cell mass = was=20 measured using CellTiter assays according to the instructions of = the=20 manufacturer (Promega, Madison, WI, http://www.promega.com/). Cell number and = viability were=20 assessed using the Guava PCA and Viacount reagent according to=20 instructions (Guava Technologies, Hayward, CA, http://www.guavatechnologies.com/). All = experiments were=20 performed in triplicate.

    -Secretase Inhibitor Synthesis

    The potent -secretase inhibitor=20 [11-endo]-N-(5,6,7,8,9,10-hexahydro-6,9-methanobenzo[a]=20 [8]annulen-11-yl)-thiophene-2-sulfonamide was listed as compound = 18 in the=20 recent report by Lewis et al. [[43]]=20 describing its synthesis and testing, and we refer to it as = GSI-18. It was=20 synthesized as previously described [[43]],=20 and its identity and quality were confirmed by nuclear magnetic = resonance=20 and mass spectral analysis. The biological activity of this = inhibitor=20 against -secretase was confirmed using in vitro and cell-based = assays as=20 previously described [[44]].=20 MRK-003 was generously provided by Merck Research Laboratories = (Boston,=20 MA, http://www.merck.com/)=20 [[43]].

    Immunocytochemistry

    The GBM neurosphere lines and low-passage cultures were seeded = in=20 12-well tissue culture plates overnight. After treating with 2 = M GSI-18 for 2-5 days, cells were = washed with=20 phosphate-buffered saline (PBS), spun down onto slides, and fixed = using 4%=20 paraformaldehyde solution in PBS at room temperature for 30 = minutes. Cells=20 were then permeabilized with 0.4% Triton X-100 in PBS for 5 = minutes at=20 room temperature, washed with PBS, and incubated in 5% bovine = serum=20 albumin (BSA)-PBS for 1 hour and then in 1:1,000 anti-NESTIN = antibody=20 (Chemicon, Temecula, CA, http://www.chemicon.com/) or 1:500 anti-Ki-67 = (DAKO,=20 Glostrup, Denmark, http://www.dako.com/) in 1% BSA-PBS for 2 hours. = After=20 washing with PBS, a final 60-minute incubation with a 1:300 = dilution of=20 cyanin 3 (Cy3)-conjugated goat anti-mouse and Cy2-conjugated goat=20 anti-rabbit secondary antibodies (Jackson Immunoresearch = Laboratories,=20 West Grove, PA, http://www.jacksonimmuno.com/) diluted in PBS = containing=20 1% BSA was done. After washing with PBS, cells were then = counterstained=20 with 4,6-diamidino-2-phenylindole (Vector Laboratories, = Burlingame, CA,=20 http://www.vectorlabs.com/), mounted, and = visualized with=20 fluorescence microscope (Carl Zeiss, Jena, Germany, http://www.zeiss.com/); 10=20 high-power fields were photographed and counted. For = immunocytochemical=20 evaluation of intracranial xenografts, frozen sections of = cryoprotected=20 brains were stained with anti-human antibody (1:100; Millipore, = Billerica,=20 MA, http://www.millipore.com/).

    Flow Cytometric=20 Analyses

    Flow cytometric analysis of CD133 was done with antibodies from = Miltenyi Biotec (Auburn, CA, http://www.miltenyibiotec.com/) according to the = instructions of the manufacturer using a FACSCalibur (Becton = Dickinson,=20 San Jose, CA, http://www.bd.com/) with CELLQuest Version 3.3 = software=20 (Becton Dickinson, San Jose, CA, http://www.bd.com/) [[42]].=20 In brief, cells were blocked in Fc receptor blocking reagent and = incubated=20 with CD133/1 (AC133)-phycoerythrin antibody (Miltenyi Biotec) for = 10=20 minutes in the dark at 4=B0C. Then cells were washed and = resuspended in 500=20 L buffer (PBS containing 0.5% BSA and = 2 mM EDTA).=20 Cells expressing levels of CD133 higher than those seen in = immunoglobulin=20 G (IgG) controls were considered positive. Experiments measuring = CD133=20 were repeated two to three times for each line and error bars = represent=20 the mean and SE.

    Real-Time Reverse=20 Transcription-Polymerase Chain Reaction

    Quantitative reverse transcription-polymerase chain reaction = (RT-PCR)=20 for NOTCH1, NOTCH2, and HES1 was done as previously described [[34][45]],=20 with all reactions normalized to actin (Applied Biosystems, Foster = City,=20 CA, http://www.appliedbiosystems.com/). Commercially = available=20 Assay on Demand TaqMan (Applied Biosystems, Foster City, CA, http://www.appliedbiosystems.com/) primers and = probes were=20 used to measure mRNA for CD133, NESTIN, BMI1, OLIG2, HES5, and = HEY1. Each=20 quantitative RT-PCR reaction was done in triplicate and error bars = represent SE.

    Protein = Analysis

    Western blots contained 20 g of protein per lane on a 10% = Tris-glycine=20 SDS-polyacrylamide gel electrophoresis gel (Invitrogen, Carlsbad, = CA, http://www.invitrogen.com/) and were = electrophoresed for=20 several hours in 1 =D7 Tris-glycine (TG)-SDS buffer (Amresco, = Solon, OH, http://www.amresco-inc.com/). Proteins were = transferred to=20 0.45-m Optitran nitrocellulose (Schleicher = &=20 Schuell, Keene, NH, http://www.schleicher-schuell.com/) in 1=D7 = Tris-glycine=20 buffer (Amresco). Blots were blocked in PBS containing 5% nonfat = dry milk=20 powder and incubated overnight at 4=B0C with antibodies directed = against=20 HES1 (1:2,000; kind gift of Dr. Tetsuo Sudo, Toray Industries, = Tebiro,=20 Kamakura, Japan, http://www.toray.com/), cleaved caspase-3 = (Asp175)=20 (1:1,000; Cell Signaling Technology, Beverly, MA, http://www.cellsignal.com/), phosphorylated Akt = (Ser473)=20 and total Akt (1:1,000; Cell Signaling Technology), phospho-STAT3=20 (pSer727) and total STAT3 (1:1,000; Affinity BioReagents, Golden, = CO, http://www.bioreagents.com/), or=20 glyceraldehyde-3-phosphate dehydrogenase (1:20,000; Research = Diagnostics,=20 Flanders, NJ, http://www.researchd.com/). Blots were then = washed several=20 times with PBS containing 0.1% Tween 20 and incubated in=20 peroxidase-conjugated IgG diluted 1:2,000 in blocking solution. = After=20 washing several times in PBS with 0.1% Tween 20, blots were = developed with=20 enhanced chemiluminescence reagent (Pierce, Rockford, IL, http://www.piercenet.com/) and exposed to = film.

    Flank and = Intracranial=20 Xenograft Tumors

    For flank xenograft tumors, 5 =D7 105 viable GBM = neurosphere=20 cells were mixed 1:1 with Matrigel (BD Biosciences, San Jose, CA, = http://www.bdbiosciences.com/) and a total = volume of 200=20 l was injected s.c. into each flank = of=20 ether-anesthetized 4-week-old female nude mice. Measurements of = tumor size=20 in the long and short axes were made weekly and estimates of = volume were=20 made using the following formula: width2 =D7 length =D7 = 0.52 [[46]].=20 For intracranial xenografts, injection guide holes were produced = in=20 anesthetized animals by twirling an 18-gauge beveled needle to = provide=20 access to the intracranial space. Fifty thousand viable cells in 2 = l of PBS were used for intracranial = injection=20 into the right striatum (0.2 L/minute) by stereotactic injection = through a=20 glass pipette connected to a Hamilton syringe. The following = coordinates=20 were used: antero-posterior =3D 0; medio-lateral =3D 2.5 mm; = dorso-ventral =3D=20 3.5 mm [[9]].=20 The skin was sutured following injection. Fifteen days after = initial=20 injection, the NOTCH inhibitor was delivered using polymer-based = beads=20 soaked with GSI-18 into the tumor beds as described previously = [[47]].=20 DMSO-soaked beads were used as controls. Tumor growth was = monitored by=20 magnetic resonance imaging (MRI) scanning at 6 weeks after GSI = treatment,=20 but survival was used as the primary endpoint in these studies. In = vivo=20 MRI studies were performed on a horizontal 9.4 T MR scanner = (Bruker=20 Biospin, Billerica, MA, http://www.bruker-biospin.com/) with a = triple-axis=20 gradient and an animal imaging probe. The scanner was also = equipped with a=20 physiological monitoring system (EKG, respiration, and body = temperature).=20 Mice were anesthetized with isoflurane (1%) together with oxygen = and air=20 at 1:3 ratio via a vaporizer and a facial mask. We used a 40-mm = diameter=20 birdcage coil for the radio frequency transmitter and receiver.=20 Temperature was maintained by a heating block built into the = gradient=20 system. Respiration was monitored throughout the entire scan and = was used=20 to synchronize the start of each acquisition. Multiple slice = T2-weighted=20 MRIs were acquired with the following parameters: fast spin echo = sequence=20 with an echo train length of 4, echo time =3D 35 ms, repetition = time =3D 3,000=20 ms, imaging resolution =3D 0.1 mm =D7 0.1 mm per pixel, slice = thickness =3D 0.3=20 mm, total imaging time of 30 minutes with respiratory = triggering.

    Statistical = analysis

    Statistical analyses were done using GraphPad Prism 4 (GraphPad = Software, http://www.graphpad.com/). Data graphed with = error bars=20 represent mean and SE from experiments done in triplicate unless = otherwise=20 noted. A two-sided Student's t test was used to determine = the=20 significance of any differences.

    RESULTS
    3DRESULTS=20 3D""=20

    GBM Neurospheres = Are Sensitive=20 to NOTCH Pathway Blockade In Vitro

    We initially studied well-characterized HSR-GBM1 neurosphere = cultures=20 (Fig. 1A),=20 which were derived from a single tumor [[9]]=20 and grown for several years as three independent sublines. The = stem cell=20 marker CD133 is expressed in these GBM-derived neurospheres, as = detected=20 using flow cytometry, in approximately 70% of the tumor cells = (Fig. 1A).=20 This line was previously shown to express markers of NOTCH = activity at the=20 time it was established [[9]],=20 and we confirmed protein expression of the NOTCH pathway target=20 HES1 in the tumor neurospheres (Fig. 1B).=20

    Figure 1. GBM = neurospheres are=20 sensitive to NOTCH pathway blockade in vitro. (A): = The GBM=20 neurosphere line HSR-GBM1 grows as large, multicellular = spheres. The=20 majority (70%) of neurosphere cells express CD133. = (B): Upon=20 GSI-18 treatment, protein levels of HES1, a NOTCH pathway = target=20 gene, were reduced after 48 hours of GSI-18 treatment of = HSR-GBM1=20 neurospheres. (C, D): Upon GSI-18 treatment, = mRNA=20 expression of NOTCH target gene HES5 and protein levels of = cleaved=20 active NOTCH1 intracellular domain are also reduced. = (E):=20 GSI-18 reduced total cell mass in a dose-dependent fashion = in all=20 three HSR-GBM1 sublines (*, p < .05). (F): = There is=20 no significant change in normal human astrocyte growth when = treated=20 with GSI-18. (G): Growth of a second GBM neurosphere = line,=20 HSR-GBM2, was also inhibited by GSI-18. (H): HES1 = mRNA levels=20 fell into a similar range in primary GBM and tumor-derived=20 neurospheres grown in culture. (I): A second -secretase inhibitor, MRK-003, could also = suppress GBM=20 neurosphere growth. (J): MRK003 showed potency = similar to=20 GSI-18 in terms of its ability to lower HES1 mRNA levels in = tumor=20 neurospheres. (K, L): Low-passage neurospheres = GBM-SC,=20 GBM-KK, and GBM-DM express the NOTCH target gene HEY1 and = had their=20 growth inhibited by NOTCH blockade. Abbreviations: DMSO, = dimethyl=20 sulfoxide; GAPDH, glyceraldehyde-3-phosphate dehydrogenase; = GBM,=20 glioblastoma; GSI-18, -secretase inhibitor 18.
    [Normal View 76K | = Magnified View=20 228K]

    NOTCH pathway blockade for 48 hours using 2 M -secretase inhibitor (GSI-18) decreased levels of HES1 = protein in=20 the HSR-GBM1 neurospheres by more than 80% (quantified by = densitometry on=20 the Western blot in Fig. 1B),=20 and no HES1 was detected when 10 M or higher concentrations of the = drug were used=20 (Fig. 1B).=20 Because HES1 can be regulated by multiple pathways, we also = examined two other markers of NOTCH signaling, HES5 = expression and=20 formation of the activated NOTCH1 intracellular domain. = Both were=20 decreased in a dose-dependent fashion by GSI-18 (Fig. 1C,=20 1D),=20 providing additional confirmation of baseline NOTCH activity and=20 pharmacologic inhibition of the pathway in tumor neurospheres. = GSI-18=20 reduced growth of all three HSR-GBM1 sublines in vitro in a = dose-dependent=20 fashion (Fig. 1E).=20 In contrast, the growth of normal human astrocytes was not = significantly=20 affected by the same level of GSI-18 (Fig. 1F),=20 indicating that this agent does not nonselectively inhibit = proliferation=20 of astrocytic cells. Two slower-growing established GBM = neurosphere lines=20 (HSR-GBM2 and HSR-GBM3) were also inhibited in a dose-dependent = fashion by=20 GSI-18 (Fig. 1G=20 and data not shown). The fact that HES1 mRNA in these GBM=20 neurosphere lines was expressed at levels within the range seen in = 24=20 snap-frozen GBM samples suggests they had biologically relevant = levels of=20 NOTCH activity (Fig. 1H).

    To help exclude the possibility that the effects of GSI-18 were = nonspecific, we also used another -secretase inhibitor with a significantly different = structure,=20 MRK-003. This second compound also suppressed HSR-GBM1 growth and = HES1=20 levels in a dose-dependent fashion (Fig. 1I,=20 1J).=20 Finally, we analyzed the growth of GBM neurospheres derived from = four=20 different tumors and grown for relatively few passages (6-20). = High NOTCH=20 activity in these cultures was confirmed by NOTCH target gene = expression=20 (Fig. 1K).=20 Growth of all four lines (SC, DM, KO, and TZ) was inhibited 70% or = more by=20 10 M MRK-003, whereas three of the lines = (SC, KO,=20 and TZ) showed a modest growth suppression by 2 M MRK-003 (Fig. 1L=20 and data not shown). These studies indicate that both = long-established and=20 lower-passage GBM-derived neurospheres require ongoing NOTCH = signaling for=20 growth in culture.

    Activation of = NOTCH2 in GBM=20 Neurospheres Increases Their Growth In Vitro

    To examine whether activation of NOTCH signaling in GBM = neurospheres is=20 sufficient to increase tumor growth, we introduced a truncated,=20 constitutively active form of the NOTCH2 receptor (NICD2) into GBM = neurosphere lines using either stable transfection or transient = expression=20 following infection by adenovirus we generated previously [[34]].=20 We used NICD2 in these experiments as NOTCH2 receptor protein has = been=20 shown to be highly expressed in glioblastoma [[48]].=20 Following NICD2 transfection and selection, HSR-GBM1 cells had=20 significantly increased mRNA levels of NOTCH2 and the = pathway=20 target HES1 (Fig. 2A-2C).=20 The elevated NOTCH2 signaling following transfection of NICD2 was=20 associated with an approximately twofold increase in growth over 5 = days in=20 culture (Fig. 2D).=20 Transient viral delivery of NICD2 also significantly increased the = growth=20 rate of GBM-derived neurospheres, albeit to a lesser degree (Fig. = 2E).=20 This less robust induction of growth may reflect the fact that = only=20 60%-80% of cells are infected by virus containing NICD2. These = findings=20 suggest that activation of NOTCH2 is sufficient to promote = GBM=20 neurosphere growth in vitro.=20

    Figure 2. Activation = of NOTCH2=20 in glioblastoma (GBM) neurospheres increases tumor growth in = vitro.=20 (A): When the active form of NOTCH2 receptor = (NICD2)=20 was stably transfected into HSR-GBM1a cells, receptor mRNA = levels=20 were elevated. (B): NICD2 protein expression was also = increased in NICD2 stably transfected HSR-GBM1 neurospheres = detected=20 by Western blot. GAPDH was used as protein loading control.=20 (C): The NOTCH target gene HES1 was also induced in=20 NICD2-transfected GBM neurospheres. (D): Cell growth = was=20 significantly increased in NICD2-transfected cells. = (E):=20 Transient expression of NICD2 in HSR-GBM1 neurospheres using = adenovirus vectors also increased cell growth over 2 days.=20 Abbreviations: Ad, Adenovirus; Beta Gal, -galactosidase; GAPDH,=20 glyceraldehyde-3-phosphate dehydrogenase; NICD2, NOTCH2=20 intracellular domain.
    [Normal View 32K | = Magnified View=20 89K]

    NOTCH Blockade = Depletes=20 CD133-Positive Cells and Inhibits Neurosphere = Clonogenicity

    We have previously shown that CD133 marks cells with=20 xenograft-initiating potential [[42][49]],=20 and that NOTCH blockade can deplete stem-like cells in = medulloblastoma=20 cultures [[42]].=20 In HSR-GBM1 neurospheres, we found an approximately twofold = increase in=20 the clonogenic potential of CD133-positive cells compared with = negative=20 ones (C.G.E. and E. Bar, unpublished data). We examined CD133 mRNA = levels=20 in HSR-GBM1 cells after NOTCH pathway blockade by GSI-18, and = found that=20 this stem cell marker was almost completely suppressed by 3-M levels of the -secretase inhibitor after 48 hours (Fig. 3A).=20 We also identified a pronounced, dose-dependent decrease in the = percentage=20 of CD133-expressing cells using flow cytometric analysis, = suggesting that=20 the cancer stem cell fraction within the tumor had been depleted = (Fig. 3B).=20 Given the evidence from two recent studies showing that the = CD133-negative=20 GBM population also forms xenografts in nude mice [[50][51]],=20 suggesting that this marker may not mark all CSCs, we also = examined=20 several other stem cell markers including NESTIN, BMI1, and OLIG2. = The=20 expression of all of these was also inhibited by GSI-18 in a=20 dose-dependent fashion, indicating a broad effect on potential CSC = markers=20 (Fig. 3C-3E).=20 We next examined clonogenicity after treatment with 5 M MRK-003 to document functional loss = of=20 clonogenic CSCs. Some tumor neurospheres were still present in = culture=20 after 5 days of NOTCH inhibition (Fig. 3F,=20 upper right panel), however, when these were dissociated, and = equal=20 numbers of viable cells seeded into semisolid neurosphere media, = those=20 that had previously been exposed to NOTCH blockade could not form = colonies=20 (Fig. 3F,=20 3G).=20 Similar but less pronounced effects were seen with 2-M levels of the -secretase inhibitor (Fig. 3G).=20 These data demonstrate that NOTCH pathway blockade depletes = stem-like=20 cells in GBM, and decreases tumor clonogenic growth in vitro.=20

    Figure 3. NOTCH = blockade=20 depletes CD133-positive cells and inhibits neurosphere=20 clonogenicity. (A): CD133 mRNA levels were severely = decreased=20 following GSI-18 treatment of HSR-GBM1 cells. (B): = The=20 percentage of CD133-positive cells in these cultures was = also=20 lowered by NOTCH blockade. (C-E): mRNA levels of the = stem=20 cell markers NESTIN, BMI1, and OLIG2 were also reduced in a=20 dose-dependent fashion following GSI-18 treatment of = HSR-GBM1=20 neurospheres. (F, G): Growing HSR-GBM1 cultures = treated for 5=20 days with -secretase inhibitor can form some neurospheres = (right top=20 panel), but when equal numbers of DMSO- and MRK003-treated = cells are=20 seeded singly in methylcellulose, the latter cannot = efficiently form=20 colonies. Abbreviations: DMSO, dimethyl sulfoxide; GAPDH,=20 glyceraldehyde-3-phosphate dehydrogenase; GBM, glioblastoma; = GSI-18,=20 -secretase inhibitor 18.
    [Normal View 51K | = Magnified View=20 149K]

    NOTCH Signaling = Blockade=20 Depletes Cells Required for GBM Engraftment In Vivo

    To extend our in vitro findings, we studied the effects of = GSI-18 on in=20 vivo tumor growth. First, we treated GBM neurospheres for 2 days = with=20 either GSI-18 or vehicle (DMSO) ex vivo, long enough to remove = most=20 stem-like GBM cells based on the in vitro experiments described = above. We=20 then injected equal numbers of surviving cells, which were = demonstrated to=20 be viable by trypan blue staining and vital count assays, = subcutaneously=20 into nude mice. After monitored tumor growth for more than 3 = months, we=20 found that GSI-18 pretreatment completely inhibited the growth of=20 subcutaneous HSR-GBM1 xenografts (Fig. 4A-4D).=20 Large tumor masses formed at all eight sites (Fig. 4A,=20 4B,=20 asterisks) injected with cells in which NOTCH signaling was = continuously=20 active, including nontreated (n =3D 4) and vehicle = (DMSO)-treated=20 (n =3D 4) cells (Fig. 4A,=20 4B).=20 In contrast, mice injected with an equal number of viable cells = previously=20 treated with 2 M GSI-18 (n =3D 6) were unable = to=20 efficiently form tumors, with small lesions developing at only = three of=20 six injection sites (Fig. 4C).=20 A comparison of growing tumor volumes in mock-, DMSO-, and 2 M GSI-18-treated xenografts is shown = in Figure 4D.=20 The small xenografts that are formed from GSI-18-treated GBM cells = had a=20 microscopic appearance quite similar to those from DMSO-treated = cells=20 (Fig. 4E,=20 4F).=20 They contained abundant HES1 protein (Fig. 4G),=20 indicating that cells that survive NOTCH blockade and grow slowly = into=20 small tumors in vivo eventually re-establish expression of NOTCH = pathway=20 targets.=20

    Figure 4. NOTCH = signaling=20 blockade depletes cells required for glioblastoma (GBM) = propagation=20 in vivo. (A-C): Large xenografts formed at all eight = sites=20 (asterisk) injected with cells in which NOTCH signaling was = active,=20 including nontreated (n =3D 4, A) and vehicle=20 (DMSO)-treated (n =3D 4) cultures. Mice injected with = equal=20 number of viable cells previously treated with 2 M GSI-18 (n =3D 6) were = unable to=20 efficiently form tumors, with small lesions developing at = only three=20 of six injection sites. (D): Tumor volume was = dramatically=20 reduced in mice engrafted with 2 M GSI-18-pretreated cells = compared with=20 mice engrafted with mock or DMSO-pretreated cells. = (E-G): The=20 small subcutaneous xenografts that formed from cells = pretreated with=20 GSI-18 appeared similar to control tumors, and expressed = abundant=20 amounts of the NOTCH pathway target HES1 on = immunohistochemical=20 analysis. (H, I): Intracranial xenografts of HSR-GBM1 = cells=20 pretreated with DMSO or GSI-18 often invaded into the = subventricular=20 space (H), and occasionally formed necrotic foci with = pseudopalisading (I). (Original magnifications both = =D7100.)=20 Abbreviations: DMSO, dimethyl sulfoxide; GSI-18, -secretase inhibitor 18.
    [Normal View 142K | = Magnified View=20 642K]

    To determine whether downregulation of NOTCH signaling prior to = injection of GBM neurospheres also affects intracranial tumor = initiation,=20 we treated HSR-GBM1 with 2 M or 5 M GSI-18 for 48 hours ex vivo to = deplete CSCs,=20 then stereotactically injected 50,000 viable cells into mouse = brains. The=20 same number of cells from DMSO-treated neurospheres was used as a = positive=20 control, with four mice included per group. Mice were sacrificed = when they=20 showed neurological signs suggesting tumor growth, and survival = was used=20 as the primary endpoint for statistical analysis. We found that = animals=20 injected with GSI-18-pretreated GBM neurospheres did form = orthotopic=20 xenografts and die, but had a significantly longer survival = compared with=20 those injected with DMSO-treated tumor cells (p < .05, = log-rank=20 analysis). Microscopic examination of the xenografts that formed = from the=20 GSI-18-pretreated (Fig. 4H,=20 4I)=20 and DMSO-pretreated (not shown) GBM neurospheres failed to = identify any=20 significant differences in the size or histopathological = appearance of the=20 gliomas, which were all large and infiltrative. Interestingly, = many of the=20 tumor xenografts, both treated and nontreated, showed a = predilection for=20 invasion and growth in the subependymal region around the = ventricles (Fig.=20 4H=20 and data not shown), perhaps due to the capacity of this area to = support=20 the growth of stem and progenitor cells. In some large tumors,=20 pseudopalisading necrosis, a common feature of glioblastoma, was = also seen=20 (Fig. 4I).=20 The fact that tumor neurosphere cultures pretreated with -secretase inhibitor did form large xenografts, albeit = more=20 slowly than controls, may reflect the fact that our in vitro = analysis=20 suggests only a partial depletion of CD133-positive cells = following NOTCH=20 blockade.

    Local Delivery of = -Secretase Inhibitors Slows Growth of Intracranial GBM=20 Xenografts

    To mimic more closely the care of GBM patients, and to test the = effects=20 of longer-term delivery of -secretase inhibitor, we also treated small = intracranial tumors=20 using local delivery of the NOTCH inhibitor. We injected GBM = neurospheres=20 into mouse brains and waited for 15 days, a time frame during = which we=20 predict that small tumors should form based on prior xenograft = time-course=20 studies. Polymer-based beads soaked with GSI-18 or DMSO were then=20 surgically implanted into the tumor beds as described previously = [[47]].=20 To confirm that GSI-18 prevented GBM neurosphere propagation in = vivo due=20 to downregulation of NOTCH signaling pathway, we also injected = cells=20 stably transfected with NICD2, thereby constitutively activating = NOTCH2=20 signaling and rendering the cells insensitive to -secretase blockade. We monitored tumor growth at 6 = weeks after=20 injection by MRI scanning in representative animals, and found = that DMSO=20 (vehicle)-treated mice form large xenografts, whereas GSI-18 = treatment=20 appeared to block formation of radiographically detectable tumors = (Fig. 5A).=20 The NICD2 transfected GBM neurospheres appeared to form larger=20 intracranial xenografts, and were not sensitive to GSI-18 = treatment (Fig.=20 5A).=20 When we compared the survival of the various groups, mice treated = with=20 GSI-18 also had a significantly prolonged survival compared with=20 DMSO-treated mice, whereas animals bearing NICD2 xenografts had = shorter=20 survival and were not sensitive to the GSI-18 treatment (Fig. 5B).=20 Microscopic examination of brains from treated animals revealed = implanted=20 polymer beads and adjacent injection tracts but no recognizable = surviving=20 tumor cells in the multiple sections examined (Fig. 5C).=20 Immunocytochemical analysis of intervening sections with = anti-human=20 antibodies that can recognize xenografted cells also failed to = identify=20 glioma (Fig. 5D,=20 5E).=20 Thus significant tumor growth is blocked in most animals following = local=20 delivery of -secretase inhibitor.=20

    Figure 5. Local GSI = treatment=20 prolongs survival of mice bearing intracranial GBM = xenografts.=20 (A): Wild-type or NICD2 stably transfected GBM = neurospheres=20 were implanted into mouse brain, and GSI-18- or DMSO-soaked = beads=20 were injected into the tumor beds 2 weeks later. Tumor = growth was=20 monitored by MRI in selected animals, and large xenografts = formed in=20 DMSO- but not GSI-18-treated mice. As expected, constitutive = NICD2=20 expression rendered xenografts insensitive to GSI-18. = (B):=20 GSI-18-treated mice have significantly prolonged survival = compared=20 with DMSO-treated mice, whereas expression of activated = NOTCH2=20 promoted more rapid intracranial tumor growth. (C-E): = Microscopic examination of a GSI-18-treated mouse brain = stained with=20 H&E reveals a round polymer bead (arrow) with a pale = linear=20 injection tract leading upward (arrowhead), but no tumor = mass on=20 this or additional step sections (original magnification = =D7100).=20 Immunocytochemical stains using an antibody specific for = human=20 nuclei confirms the absence of viable tumor cells (D, = E).=20 (Original magnification =D7100.) Abbreviations: DAPI, 4,6-diamidino-2-phenylindole; DMSO, dimethyl = sulfoxide;=20 GAPDH, glyceraldehyde-3-phosphate dehydrogenase; GBM, = glioblastoma;=20 GSI-18, -secretase inhibitor 18; Hu, human nuclei; MRI, = magnetic=20 resonance imaging; NICD2, NOTCH2 intracellular = domain.
    [Normal View 68K | = Magnified View=20 274K]

    NOTCH Blockade = Selectively=20 Reduces Proliferation of Stem-Like GBM Cells

    To examine the mechanism by which NOTCH regulates stem-like = cells in=20 GBM, we used double immunolabeling to assess proliferation in = cells=20 expressing the stem/progenitor markers. Because it has been shown = that=20 NOTCH regulates NESTIN expression in GBM [[52]],=20 raising the possibility that population changes could be due to = direct=20 effects of NOTCH blockade on NESTIN levels rather than loss of = stem-like=20 cells, we used both NESTIN and CD133 as markers. Consistent with = the=20 concept that ongoing NOTCH signaling is required in poorly = differentiated=20 cells, we found that pathway blockade by GSI-18 significantly = reduced the=20 percentage of cells expressing high levels of NESTIN or CD133 in = GBM=20 neurospheres (Fig. 6A,=20 6B).=20 The overall Ki-67-positive proliferating fraction of cells was = also=20 significantly reduced (Fig. 6B).=20 Importantly, proliferation rates were most dramatically lowered by = GSI-18=20 in the NESTIN- or CD133-bright population, whereas NESTIN- or = CD133-dim=20 cells showed a far lesser reduction in cycling cells (Fig. 6C,=20 6D).=20 This suggests that NOTCH inhibition depletes the stem-like tumor=20 subpopulation at least in part due to reduced proliferation. = Although it=20 is not entirely clear how the proliferation rate of stem-like = glioma cells=20 might affect tumor growth, it has recently been shown that the = presence of=20 a high percentage of glioblastoma cells coexpressing CD133 and = Ki-67 is=20 associated with worse clinical outcomes [[53]].=20

    Figure 6. NOTCH = signaling=20 regulates proliferation of poorly differentiated GBM cells.=20 (A): NOTCH pathway blockade by GSI-18 reduces the = NESTIN- or=20 CD133-positive population in neurosphere lines, as assessed = by=20 double immunolabeling (upper panel: NESTIN in green; Ki-67, = red;=20 DAPI, blue; lower panel: CD133 in red; Ki-67, green; DAPI, = blue).=20 (B): Both NESTIN- and Ki-67-positive populations were = significantly reduced. (C, D): Ki-67 proliferation = indices=20 were significantly lowered by GSI-18 in the NESTIN- or=20 CD133-positive cell population, whereas NESTIN- or = CD133-negative=20 cells showed a lesser reduction (p value for t = tests:=20 *, p < .05, **, p < .01). Abbreviations: = DAPI,=20 4,6-diamidino-2-phenylindole; DMSO, dimethyl = sulfoxide;=20 GSI-18, -secretase inhibitor 18.
    [Normal View 73K | = Magnified View=20 238K]

    NOTCH Signaling = Regulates=20 Phosphorylation of Akt and STAT3 in GBM Neurospheres

    To address the possibility that NOTCH blockade induces cell = death in=20 GBM neurosphere cells, as well as decreasing proliferation, we = examined=20 cleavage of caspase-3 following GSI-18 treatment. An increase in = the=20 proapoptotic cleaved form of this protein was identified in = cultures=20 treated with 0.4 M and higher levels of the NOTCH = pathway=20 inhibitor, suggesting that cell death was induced by NOTCH = blockade (Fig.=20 7).=20 We also wanted to identify the downstream targets that might = mediate the=20 apoptotic effects of NOTCH pathway blockade in GBM neurospheres. = AKT and=20 p53 are two key proteins that regulate apoptosis in many type of = cancers=20 [[54]].=20 We therefore examined whether protein levels or phosphorylation = states of=20 p53 and AKT were changed in GSI-18-treated GBM neurospheres. We = did not=20 observe p53 alteration in GSI-18-treated GBM cells (data not = shown).=20 However, 2 M GSI-18 totally blocked AKT = phosphorylation=20 (pAKT-Ser473) in the GBM neurosphere line HSR-GBM1 (Fig. 7).=20 These results suggest that phosphorylation and activation of AKT = promoted=20 by NOTCH may mediate survival of GBM CSCs. Recently, it has been = shown=20 that NOTCH signaling can regulate stem cell numbers both in vitro = and in=20 vivo through phosphorylation of STAT3 (pS727) [[55]].=20 To determine whether STAT3 might also be affected by NOTCH = inhibition in=20 brain tumors, we examined pSTAT3 protein changes in GSI-18-treated = HSR-GBM1 neurospheres. We found that phosphorylation of STAT3 was=20 significantly reduced by NOTCH pathway blockade in a = dose-dependent=20 fashion, whereas total STAT3 was essentially unchanged in = GSI-18-treated=20 HSR-GBM1 neurospheres (Fig. 7).=20 These results suggested that NOTCH signaling may regulate GBM CSC = survival=20 through phosphorylation of AKT and STAT3.=20

    Figure 7. NOTCH = signaling=20 blockade promotes caspase-3 cleavage and reduces = phosphorylation of=20 AKT and STAT3. (A): Loss of AKT and STAT3 = phosphorylation and=20 activation of caspase-3 cleavage is induced by 2 M or higher levels of GSI-18, = whereas=20 overall levels of AKT and STAT3 are unchanged in GBM = neurosphere=20 cells. GAPDH is used as a loading control. These changes = were=20 quantitated using densitometry in (B-D). = Abbreviations: DMSO,=20 dimethyl sulfoxide; GAPDH, glyceraldehyde-3-phosphate = dehydrogenase;=20 GBM, glioblastoma; GSI-18, -secretase inhibitor 18.
    [Normal View 41K | = Magnified View=20 109K]

    DISCUSSION
    3D""=20 3DDISCUSSION=20

    A number of groups analyzing GBM have identified molecular = changes in=20 NOTCH pathway members, implicating activation of this = developmentally=20 important signaling cascade in the pathobiology of malignant brain = tumors=20 [[9][12][32]].=20 In the current study, we used neurospheres derived from human GBM=20 specimens to examine the effects of NOTCH pathway blockade. Such=20 neurosphere lines represent an improved model system, as when = injected=20 orthotopically into immunocompromised mice they recapitulate the=20 pathobiology of GBM more closely than traditional adherent cell = lines=20 maintained in high serum. Both established and lower-passage = neurosphere=20 cultures were used. We found that NOTCH pathway blockade by either = of two=20 structurally distinct -secretase inhibitors slows tumor growth in all GBM = neurosphere=20 cultures tested, whereas overexpression of a constitutively active = form of=20 NOTCH2 increases tumor growth. In addition, we demonstrate that = NOTCH=20 pathway blockade inhibits GBM neurosphere engraftment in vivo, and = significantly prolongs the survival of mice bearing intracranial=20 xenografts. Larger preclinical studies will be needed to confirm = and=20 extend these data.

    It has been suggested that NOTCH may play a particularly = important role=20 in CSCs, a subpopulation of tumor cells that have stem-like = properties and=20 are uniquely capable of enabling engraftment and long-term growth = of=20 cancers [[5-10][32][56-58]].=20 Although a few gain-of-function studies directly support the = concept that=20 NOTCH is important in stem-like GBM cells [[52][59]],=20 to our knowledge it has not been shown that NOTCH pathway = inhibition can=20 deplete CSCs in gliomas. We found that pharmacological NOTCH = blockade=20 using -secretase inhibitors reduces the percentage of cells = expressing=20 the stem/progenitor cell markers CD133 and NESTIN in GBM = neurospheres. In=20 addition, following NOTCH blockade, surviving cells are no longer = able to=20 efficiently form colonies in vitro or engraft in vivo, consistent = with the=20 concept that a key subpopulation of cells required for efficient = tumor=20 propagation is no longer present. We believe that the depletion of = the GBM=20 subpopulation expressing stem cell markers, in conjunction with = the loss=20 of clonogenic and engraftment potential, suggests that NOTCH = blockade can=20 at least partially remove CSCs from GBM neurosphere lines.

    Gastrointestinal toxicity has been a major concern when using = GSI, with=20 NOTCH blockade inducing goblet cell differentiation in intestinal = transit=20 amplifying cells, leading to severe diarrhea [[60-63]].=20 To avoid such side effects, we used polymer-based beads to deliver = GSI-18=20 locally within the brain in our in vivo experiments. This method = is=20 conceptually similar to local delivery of the chemotherapeutic = agent=20 biodegradable carmustine via polymer wafers to the tumor bed, a = Food and=20 Drug Administration (FDA)-approved strategy currently used in GBM = patients=20 [[64-67]].=20 The fact that this local delivery of GSI-18 significantly prolongs = survival without any detectable gastrointestinal side effects in = animals=20 suggests a strategy for translating GSI-based therapies into=20 neuro-oncology clinics. One additional potential concern when = targeting=20 CSCs in the brain is that NOTCH inhibition may also deplete normal = neural=20 stem cells. However, we did not find obvious neurological defects = in the=20 animal receiving GSI over several months of observation.

    To maximize the efficacy of anti-CSC agents such as -secretase inhibitors, it will likely be necessary to = give them=20 in combination with other treatment modalities (surgery, radiation = therapy, other chemotherapy). Indeed, it has recently been shown = that=20 giving a -secretase inhibitor in conjunction with the = glucocorticoid=20 dexamethasone can potentiate its therapeutic effect and reduce = systemic=20 side effects when treating leukemia [[68]].=20 In this regard, our discovery that the reduced proliferation and = induction=20 of apoptotic markers in GBM following NOTCH blockade is associated = with=20 phosphorylation changes to AKT and STAT3 suggests additional = pathways that=20 might be synergistically targeted.

    SUMMARY
    3D""=20 3DSUMMARY=20

    In summary, our data support the concept that -secretase inhibitors can inhibit the growth of = functionally=20 heterogeneous GBM neurospheres, both in vitro and in vivo, with=20 particularly pronounced effects on poorly differentiated tumor = cells. The=20 potential therapeutic implications of targeting of stem-like cells = via=20 NOTCH inhibition extend beyond neuro-oncology, as this signaling = pathway=20 has been shown to play a role in CSCs not only in neural tumors, = but also=20 in cancers from breast, lung, pancreas, and prostate [[26][69-72]].=20 More studies will be needed to determine whether pharmacological = NOTCH=20 blockade, either alone or in conjunction with other therapies, = will be=20 effective in improving the survival of patients with GBM and other = malignant tumors.

    Disclosure of Potential Conflicts = of=20 Interest

    The authors indicate no potential conflicts of interest.

    Acknowledgements
    3D""=20 3DAcknowledgements=20

    We acknowledge grant support to X.F. from Accelerate Brain = Cancer Cure=20 Project Award, American Brain Tumor Association Translational = Grant, and=20 Voices Against Brain Cancer Research Grant, and to C.G.E. from = NIH/NS55089=20 and the Brain Tumor Funders Collaborative. We also acknowledge = Merck and=20 Merck scientists for providing MRK-003.

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