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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_01CAD4B7.1560CE90 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_01CAD4B7.1560CE90 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: 17-28
    • Top of page
    • Abstract
    • INTRODUCTION
    • MATERIALS AND METHODS
    • RESULTS
    • DISCUSSION
    • Acknowledgements
    • References
    List of Images
    Abstract=20 Only

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***************************=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 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    3D"About

    Online ISSN: 1549-4918    Print ISSN: = 1066-5099
    STEM CELLS
    Volume 28, Issue 1, 2010. Pages: 17-28

    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. -Secretase = inhibitors (GSIs) reduce cell growth and clonogenic survival of glioma = stem cells after radiation. (A): T4302 CD133+ glioma stem cells = were transfected with a Notch-responsive luciferase reporter (RBP-J reporter) premixed with = the vector directing constitutive expression of Renilla = luciferase. Cells were left unirradiated or irradiated at 3 Gy and were = then collected to measure luciferase activities at the time points as = indicated. In this and subsequent luciferase assays, firefly luciferase = activity was normalized to the internal control Renilla = luciferase activity, and the normalized firefly luciferase activity from = the sham-irradiated control group was assigned a value of 1. Data are = presented as the mean =B1 standard error. (B): T4302 CD133+ = glioma stem cells were left unirradiated or irradiated at 3 Gy. The = relative mRNA levels of Hes2 were determined by quantitative real-time = polymerase chain reaction at the time points as indicated. (C): = T4302 CD133+ cells were transfected with the RBP-J reporter and treated with = dimethyl sulfoxide (vehicle), 2 M DAPT, or 0.5 M = L685,458 overnight, and cells were then collected for luciferase assays. = (D): T4302 CD133+ cells were pretreated with 2 M DAPT or = 0.5 M L685,458 for 4 hours and irradiated as indicated. = Total viable cell numbers were then determined with a cell viability = assay for the following 5 days. The averaged cell titers on day 1 of the = control groups were assigned a value of 1. (E): T4302 CD133+ = cells were pretreated and irradiated as described in (D). Cells = were then plated to form neurospheres, and the surviving fractions were = determined as described in Materials and Methods. The mean =B1 standard error = was derived from eight replicates within a representative experiment. = (F,G): Neurosphere formation of T3359 and T4105 CD133+ cells = treat with GSIs alone, 3 Gy radiation alone, or the combination was = determined as described in (E). *P < .05 by = Student's t test; #P < .001 by = one-way ANOVA. Abbreviation: DAPT, N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester.
    '); } =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. -Secretase = inhibitors (GSIs) enhance radiation-induced cell death in glioma stem = cells. (A): T4302 CD133+ cells were pretreated with DAPT or = L685,458 for 4 hours and irradiated at 3 Gy. Percentage of dead cells = was determined with trypan blue exclusion assay at 48 hours after = radiation exposure. (B): T4302 CD133+ cells were treated with = GSIs =B1 radiation as indicated. Relative caspase 3/7 activities were = determined by normalizing caspase activities to the corresponding cell = titers. (C): T4302 CD133+ cells were treated as described in = (A).At 24 or 48 hours after radiation exposure, cells were = labeled with FITC-conjugated annexin V staining and analyzed with flow = cytometry. (D): T4302 CD133+ cells were pretreated with 2 M DAPT = =B1 radiation as indicated. Relative caspase 3/7 activities were = determined at 72 hours after radiation. (E) T4302 CD133+ cells = were treated with DAPT at indicated concentrations for 4 hours and = irradiated at 3 Gy. Relative caspase activities were determined at 48 or = 72 hours after radiation. *P < .01 by Student's t test; #P < .01 by one-way ANOVA. = Abbreviation: DAPT, N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester.
    '); } =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. Expression of the intracellular domains of Notch1 or Notch2 attenuates = the radiosensitizing effects of -secreting inhibitors (GSIs). T4302 CD133+ glioma cells = were infected with control lentivirus or lentivirus directing expression = of NICD1 or NICD2 and selected by puromycin. (A): Expression of = Notch1, Notch2, Hes2, Hes4, and Hes5 were determined by quantitative = real-time polymerase chain reaction. (B): Control cells or = cells expressing NICD2 were transfected with Notch-dependent luciferase = reporter and treated with dimethyl sulfoxide or 2 M DAPT = overnight. Notch-dependent transcription activities were determined by = luciferase reporter assays as described in Figure 1. (C): = Control cells or cells expressing NICDs were treated with DAPT =B1 = radiation as indicated. Relative caspase 3/7 activities were determined = at 48 or 72 hours after radiation. (D): Apoptotic cell death = was determined with annexin V staining as described in Figure 2. (E): = Surviving fractions were determined as described in Figure 1. *P = < .01 by Student's t test; #P < = .001 by one-way ANOVA. Abbreviations: DAPT, N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester; NICD, Notch receptor intracellular domain.
    '); } =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 pathway does not alter DNA damage response of glioma stem cells. = (A): T4302 CD133+ cells were pretreated with DAPT or L685,458 = for 24 hours before radiation. At 1 hour after radiation exposure, cells = were harvested. The levels of activating phosphorylation of Chk1 and = Chk2 were assessed by immunoblotting. (B): T4302 CD133+ cells = were treated with 2 M DAPT at 4 and 24 hours before radiation (-24 hours = or -4 hours) or at 0, 4, and 24 hours after radiation. Relative caspase = 3/7 activities were measured 3 days after radiation exposure. *P < .001 by Student's t test. Abbreviation: DAPT, N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester.
    '); } =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. Notch pathway regulates radioresistance of glioma stem cells via the = Akt pathway and Mcl-1. T4302 CD133+ cells were infected with control = lentivirus or lentivirus directing expression of NICD2. Cells were = pretreated with or without 2 M DAPT for 4 hours and exposed to 3 Gy = radiation. Irradiated cells were collected at 24 or 48 hours after = radiation, and sham-irradiated cells were collected at 48 hours after = radiation. (A, C, D): Protein levels of phospho-Akt (S473), = total Akt, PTEN, Bcl-2, Bcl-xL, Mcl-1, phospho-p53 (S15), total p53, and = PUMA were assessed by immunoblotting. Actin was blotted as the loading = control. (B): Control cells or NICD2-expressing cells were = pretreated with 10 M PI3K inhibitor, 10 M = LY294002, or 50 M Akt inhibitor III (SH-6), for 4 hours before = radiation exposure. Relative caspase 3/7 activities were determined at = 48 hours after radiation exposure. #P < .001 by = one-way ANOVA. Abbreviations: DAPT, N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglycine t-butyl ester; NICD, Notch receptor intracellular domain.
    '); } =0A= function gcaption_fig005(hwnd) { hwnd.document.write('AlphaMed Press

    '); } =0A= function display_fig005(size) { var i =3D new Array(); i[0] =3D = 'nfig005'; i[1] =3D 'mfig005'; var hwnd =3D newfigwin(); dochead(hwnd, = ''); copyright(hwnd); otherlinks(hwnd, 'fig005', size,'null'); = imagelink(hwnd, i[size]); gcaption_fig005(hwnd); caption_fig005(hwnd); = doctail(hwnd); hwnd.document.close(); hwnd.focus(); } =0A= function caption_fig006(hwnd) { hwnd.document.write('Figure 6. Notch inhibition by -secreting inhibitors (GSIs) does not significantly alter = radiation response of CD133-negative glioma cells. T3359 CD133-negative = cells were pretreated with 2 M GSI or 0.5 M = L685,458 for 4 hours and irradiated at 3 Gy or as indicated. (A): Relative caspase 3/7 activities were determined at 48 or 72 hour = after radiation as described in Figure 2. (B): = After radiation, cells in 6-well plates were cultured for 3 weeks and = then fixed and stained to count colonies. Surviving fractions were = determined as described in Materials and Methods. (C, D): The = effects of GSIs on cell death and clonogenic survival of T4302 = CD133-negative cells were determined as described above. ##P > .05 by one-way ANOVA. (E): T4302 = CD133-negative cells were treated with GSIs =B1 radiation as described = above. Levels of phospho-Akt (S473), total Akt, Mcl-1, and actin were = determined by immunoblotting.
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    '); } =0A= function display_fig006(size) { var i =3D new Array(); i[0] =3D = 'nfig006'; i[1] =3D 'mfig006'; var hwnd =3D newfigwin(); dochead(hwnd, = ''); copyright(hwnd); otherlinks(hwnd, 'fig006', size,'null'); = imagelink(hwnd, i[size]); gcaption_fig006(hwnd); caption_fig006(hwnd); = doctail(hwnd); hwnd.document.close(); hwnd.focus(); } =0A= function caption_fig007(hwnd) { hwnd.document.write('Figure 7. Knockdown of Notch1 or Notch2 increases radiosensitivity of glioma stem = cells in vitro and in vivo. T4302 CD133+ cells were infected with = lentivirus directing expression of nontargeting shRNA or shRNAs specific = to Notch1 or Notch2 (KD-Notch1 or KD-Notch2) and selected with = puromycin. (A): Relative mRNA levels of Notch1, Notch2, Hes4, = or Hes5 were determined with quantitative real-time polymerase chain = reaction. *P < .05 by Student's t test. (B): Relative caspase 3/7 activities were determined in control = cells or cells with Notch knockdown treated with or without radiation. = #P < .001 by one-way ANOVA. (C): = Clonogenic survival was compared between the control group and cells = with Notch knockdown as described in Figure 1. (D): = Representative images (=D7100) of neurospheres formed by T4302 CD133+ = cells with or without Notch knockdown 2 weeks after radiation. (E): After lentivirus infection and puromycin selection, T4302 CD133+ = cells were left unirradiated or irradiated at 3 Gy, and they were = immediately injected into both flanks of athymic nude mice at 250,000 = cells per injection. Three mice were inoculated for a total of six = injections per group. Tumor growth was monitored twice a week for 4 = weeks until diameters of the largest tumors reached approximately 20 = mm.
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    3D""=20

    STEM CELLS

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    Volume 28, Issue 1, Pages 17-28

    Published Online: 17 Nov 2009

    Copyright =A9 2010 AlphaMed Press

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    Abstract  |  References  |  Full Text: = HTML , PDF (1072k)  |  Supporting Information=20

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    3D""=20
     Cancer Stem Cells
    Notch Promotes Radioresistance of = Glioma=20 Stem Cells =A7
    Jialiang Wang 1 2=20 *=B6, Timothy P. Wakeman=20 3, Justin D. Lathia = 4,=20 Anita B. Hjelmeland 4, = Xiao-Fan=20 Wang 3, Rebekah R. = White 1 2, Jeremy N. Rich=20 4 *||, Bruce = A.=20 Sullenger 1 2=20 *
    1Department of = Surgery,Duke=20 University Medical Center, Durham, North = Carolina
    2Duke Translational Research = Institute,Duke=20 University Medical Center, Durham, North = Carolina
    3Department of Pharmacology and = Cancer Biology,=20 Duke University Medical Center, Durham, North = Carolina
    4Department of Stem Cell Biology and=20 Regenerative Medicine, Cleveland Clinic, Cleveland, = Ohio
    email: Jialiang Wang (jialiang.wang@duke.edu)=20 Jeremy N. Rich (richj@ccf.org)=20 Bruce A. Sullenger (bruce.sullenger@duke.edu)

    *Correspondence to Jialiang = Wang, Box=20 103035, 1045 MSRB II, 106 Research Drive, Durham, NC 27710

    *Correspondence to Jeremy N. = Rich, 9500=20 Euclid Avenue, NE30, Cleveland, Ohio 44195

    *Correspondence to Bruce A. = Sullenger,=20 Box 103035, 1087 MSRB II, 106 Research Drive, Durham, NC=20 27710

    Author contributions: J.W.: Conception and = design,=20 financial support, collection and/or assembly of data, data = analysis and=20 interpretation, manuscript writing, final approval of manuscript; = T.P.W.:=20 Collection and/or assembly of data; J.D.L.: Collection and/or = assembly of=20 data; A.B.H.: Collection and/or assembly of data, data analysis = and=20 interpretation; X.-F.W.: Financial support; R.R.W.: Financial = support;=20 J.N.R.: Financial support, conception and design, manuscript = writing,=20 final approval of manuscript; B.A.S.: Financial support, = conception and=20 design, manuscript writing, final approval of = manuscript.
    Disclosure of potential conflicts of interest is = found at=20 the end of this article.
    =A7First published online = in STEM=20 CELLS EXPRESS November 17, = 2009.
    =B6Telephone:=20 919-684-4822; Fax: 919-684-8492
    ||Telephone: = 216-636-0790;=20 Fax: 216-636-5454
    Telephone: 919-684-6375; Fax: 919-684-8492

    Funded by:
     Basic Research Fellowship from the American Brain = Tumor=20 Association
     Howard Hughes Medical Institute Early Career = Award
     Basic Research Fellowship from the American Brain = Tumor=20 Association
     National Service Research Award; Grant Number: NCI = F32=20 CA142159
     National Institutes of Health (NIH); Grant Number: = CA129190
     Childhood Brain Tumor Foundation
     Pediatric Brain Tumor Foundation of the United=20 States
     Accelerate Brain Cancer Cure
     Alexander and Margaret Stewart Trust
     Brain Tumor Society
     Goldhirsh Foundation
     Sidney Kimmel Foundation
     Damon Runyon Cancer Research Foundation
     NIH grants; Grant Number: NS047409, NS054276, = CA129958,=20 CA116659

    Keywords
    Notch =95 Cancer stem cells =95 Glioma =95 Radioresistance = =95=20 Radioresponse

    Abstract
    3DAbstract=20
    Radiotherapy represents the most effective nonsurgical=20 treatments for gliomas. However, gliomas are highly = radioresistant=20 and recurrence is nearly universal. Results from our = laboratory and=20 other groups suggest that cancer stem cells contribute to=20 radioresistance in gliomas and breast cancers. The Notch = pathway is=20 critically implicated in stem cell fate determination and = cancer. In=20 this study, we show that inhibition of Notch pathway with = -secretase inhibitors (GSIs) renders the glioma = stem cells=20 more sensitive to radiation at clinically relevant doses. = GSIs=20 enhance radiation-induced cell death and impair clonogenic = survival=20 of glioma stem cells but not non-stem glioma cells. = Expression of=20 the constitutively active intracellular domains of Notch1 or = Notch2=20 protect glioma stem cells against radiation. Notch = inhibition with=20 GSIs does not alter the DNA damage response of glioma stem = cells=20 after radiation but rather reduces Akt activity and Mcl-1 = levels.=20 Finally, knockdown of Notch1 or Notch2 sensitizes glioma = stem cells=20 to radiation and impairs xenograft tumor formation. Taken = together,=20 our results suggest a critical role of Notch signaling to = regulate=20 radioresistance of glioma stem cells. Inhibition of Notch = signaling=20 holds promise to improve the efficiency of current = radiotherapy in=20 glioma treatment. STEM CELLS = 2010;28:17-28

    Received: 17 August 2009; Accepted: 6 November 2009

    Digital Object = Identifier=20 = (DOI)

    10.1002/stem.261  About=20 DOI

    Article=20 Text

    INTRODUCTION
    3DINTRODUCTION=20

    Malignant gliomas are among the most devastating malignancies. = Despite=20 recent advances in therapy, treatment of malignant gliomas remains = palliative [[1-3]].=20 Maximal surgical resection of the tumor mass followed by = radiotherapy and=20 chemotherapy is the standard of care. Gliomas often respond to=20 radiotherapy, but subsequent recurrence is almost inevitable, = suggesting=20 insufficient killing of tumorigenic cells [[2][4]].=20 Recently, cancer cells with stem cell-like properties have been = described=20 in a wide range of human tumors. The cancer stem cell model = suggests a=20 hierarchical organization of tumors that a subpopulation of tumor = cells at=20 the apex drives and maintains human tumors [[5]].=20 Cancer stem cells of brain tumors have been prospectively enriched = by=20 selection of the CD133 (Prominin-1) cell surface marker [[6-11]],=20 although some brain tumors may not express CD133, and hence = CD133-negative=20 cells may still have characteristics of cancer stem cells [[12][13]].=20 Despite expression of certain neural and other stem cell markers = (such as=20 CD133, Musashi-1, Nestin, Sox2, and Olig2) by brain tumor stem = cells, the=20 definition of cancer stem cells remains functional, requiring = sustained=20 self-renewal and tumor propagation. Our laboratory previously = demonstrated=20 that glioma stem cells were more resistant to radiation than = matched=20 non-stem glioma cells due to preferential activation of the DNA = damage=20 response pathway [[7]].=20 Other groups also reported that the cancer stem cells of breast = cancers=20 were relatively resistant to radiation, potentially due to lower = levels of=20 reactive oxygen species found in cancer stem cells [[14]].

    The emerging role of cancer stem cells in tumor response to=20 radiotherapy urges investigation on molecular mechanisms = underlying=20 radioresistance of these cells. One of these candidates is the = Notch=20 pathway. Notch mediates short-range cellular communication through = interaction with ligands presented on neighboring cells [[15]].=20 The instrumental roles of Notch in regulation of self-renewal and = cell=20 fate determination in normal stem cells have been well established = [[16]].=20 In mammals, Notch signals through four Notch receptors (Notch 1-4) = and=20 five ligands (Jagged-1 and -2, and Delta-like-1, -3, and -4) [[17]].=20 Activation of Notch involves sequential proteolytic cleavages that = eventually lead to release and nuclear translocation of the = intracellular=20 domains of Notch receptors (NICDs) and subsequent activation of=20 Notch-dependent transcription. The -secretase complex, which mediates the last proteolytic = step for=20 release of the NICDs, is essentially required for Notch activation = [[18][19]].=20 Inhibitors of -secretase (GSIs) are widely used to block Notch = signaling in=20 vitro and in vivo.

    The Notch signaling pathway is essentially involved in the = maintenance=20 of a variety of adult stem cells, such as breast [[20][21]],=20 intestinal [[22][23]],=20 and neural stem cells [[24-26]],=20 through promoting self-renewal and repressing differentiation. = Aberrant=20 Notch activity is found in a wide range of human tumors, such as = leukemia=20 [[27]],=20 breast cancer [[28][29]],=20 and glioma [[30]].=20 Taking into account the instrumental functions of Notch in both = stem cells=20 and cancer biology, it is not surprising to find that Notch plays = an=20 important role in cancer stem cells [[31]].=20 Notch inhibition impairs the self-renewal capacity of=20 mammosphere-initiating cells derived from in situ ductal carcinoma = [[32]]=20 and reduces colony formation by leukemic stem cells [[33]].=20 Activation of Notch through expression of NICD1 promotes growth = and=20 neurosphere formation of the SHG-44 glioma cell line [[34]].=20 Notch inhibition by GSI-18 induces apoptosis and differentiation = in CD133+=20 stem-like cells enriched from medulloblastoma cell lines and = impairs the=20 tumorigenic capacity of these cells [[35]].=20 In this study, we questioned the potential role of Notch in = regulating=20 glioma stem cell radioresistance using pharmacological and genetic = targeting strategies with an additional focus on potential = molecular=20 mechanisms and present data, which suggest that integrated Notch = signaling=20 is critically involved in radioresistance of glioma stem = cells.

    MATERIALS AND=20 METHODS
    3D"MATERIALS

    Cell Culture and = Enrichment of=20 Glioma Stem Cells

    Matched cultures enriched or depleted for glioma stem cells = were=20 isolated from human surgical specimens (T3359, T3691, T4105, = T4302, and=20 T4597) that were immediately implanted in immunocompromised mice = (athymic=20 BALB/c nu/nu) according to a method that has been described in our = previous studies and those of others to preserve cancer stem cells = in=20 glioma models [[7][36-40]].=20 Briefly, tumors were dissected, washed in Earle's balanced salt = solution,=20 digested with papain (Worthington Biochemical, Lakewood, NJ, http://www.worthington-biochem.com/) and = filtered through=20 70-m cell strainers to remove tissue = pieces. Red=20 blood cells were lysed in diluted phosphate-buffered saline = solution=20 (0.25=D7). Dissociated cells were then cultured overnight in stem = cell media=20 (neurobasal media supplemented with B27, epidermal growth factor, = and=20 basic fibroblast growth factor at 20 ng/ml; Invitrogen, Carlsbad, = CA, http://www.invitrogen.com/) before cell sorting = for=20 recovery of cellular surface antigens. The CD133-negative and = CD133+=20 fractions were separated by magnetic sorting using the CD133 = Microbead kit=20 (Miltenyi Biotec, Bergisch Gladbach, Germany, http://www.miltenyibiotec.com/). CD133-negative = cells were=20 maintained in Dulbecco's modified Eagle's medium supplemented with = 10%=20 fetal bovine serum (Invitrogen) but were cultured in stem cell = media at=20 least 24 hours prior to experiments to control differences in cell = media.=20 The cancer stem cell properties of the CD133+ cells were confirmed = with=20 fluorescent in situ hybridization, serial neurosphere assays, and=20 xenotransplantation assays, but cultures depleted of cancer stem = cells did=20 not initiate tumors ([[7][36-40]]=20 and data not shown).

    Antibodies

    The antibodies purchased from Cell Signaling Technology = (Beverly, MA,=20 http://www.cellsignal.com/) include = phosphor-S296-Chk1=20 (#2349), total Chk1 (#2345), phosphor-T68-Chk2 (#2661), total Chk2 = (#2662), phospho-S473 Akt (#9271), total Akt (#2920), PTEN = (#9556),=20 phospho-S15-p53 (#9286), Bcl-2 (#2870), Bcl-xL (#2764), Mcl-1 = (#4572), and=20 Puma (#4976). Other antibodies used in this study are p53 antibody = (#sc-126, Santa Cruz Biotechnology Inc., Santa Cruz, CA, http://www.scbt.com/) and=20 actin antibody (Millipore, Billerica, MA, http://www.millipore.com/).

    Radiation

    Cells were irradiated as a single-cell suspension for CD133+ = cells or=20 as monolayer culture for CD133-negative cells using an AGFA X-RAD = 320=20 irradiation system at 0.85 Gy/min.

    Growth Curve, = Caspase 3/7=20 Assay, and Clonogenic Survival Assay

    Cells were aliquoted into a 96-well plate at 3,000 cells per = well in=20 triplicate immediately after treatments. Cell number was measured = daily=20 for 5 days using the CellTiter-Glo assay kit (Promega, Madison, = WI, http://www.promega.com/).=20 Averaged cell titer of the sham-treated control group on day 1 was = assigned a value of 1. All other relative cell titers were = normalized=20 accordingly.

    The caspase 3/7 activities at 48 or 72 hours after radiation = were=20 measured using the Caspase-Glo 3/7 assay (Promega) according to = the=20 manufacturer's instructions. The caspase activities were = normalized to the=20 corresponding cell titers to generate the relative caspase 3/7=20 activities.

    To measure clonogenic survival, CD133+ glioma cells were plated = in=20 24-well plates at 50 cells per well for the sham-irradiated = groups, 100=20 cells per well for cells irradiated at 1 Gy, or 500 cells per well = for=20 cells irradiated at 2 or 3 Gy. Eight wells were plated for each = group. At=20 14 days after plating, neurospheres containing more than 50 cells = were=20 scored. Alternatively, CD133-negative glioma cells were plated in = 6-well=20 tissue culture plates in triplicates at 100 cells per well for the = sham-irradiated groups, 200 cells per well for cells irradiated at = 1 Gy,=20 or 500 cells per well for cells irradiated at 2 or 3 Gy. At 21 = days after=20 radiation, cells were fixed and stained with 0.5% crystal violet. = Colonies=20 consisting of more than 50 cells were scored. To determine the = surviving=20 fractions, the number of neurospheres at each radiation dose was=20 normalized to that of the corresponding sham-irradiated group.

    RESULTS
    3DRESULTS=20

    GSIs Impair Cell = Growth and=20 Clonogenic Survival of Glioma Stem Cells in Combination with=20 Radiation

    Accumulating evidence suggests that the Notch pathway plays = important=20 roles in a wide range of human tumors as well as cancer stem cells = [[31][41]].=20 Because a recent study reported that radiation induced transient=20 upregulation of Notch pathway components in endothelial cells [[42]],=20 we investigated whether Notch activity is involved in radiation = response=20 of glioma stem cells. Short-term cultures enriched for glioma stem = cells=20 were isolated from surgical biopsy specimens and amplified because = direct=20 xenografts that have been demonstrated to maintain a cancer stem = cell=20 phenotype [[39]].=20 The stem cell-like properties of the CD133+ glioma cells = specifically used=20 in this study, including sustained self-renewal, expression of = stem cell=20 markers, multilineage differentiation, and the ability to = recapitulate the=20 original tumors in serial xenotransplantation assays, have been=20 demonstrated in our previous publications [[6][7][36-38][40]].=20 In T3359, T4105, and T4302 CD133+ glioma cells, radiation induced = modest=20 transcription activation from a Notch-responsive luciferase = reporter=20 (RBP-J=20 reporter; SABiosciences, Frederick, MD, http://www.sabiosciences.com/) but not from a = control=20 reporter (SABiosciences) (Fig. 1A,=20 and supporting information Figure 1A-1C). Expression of Notch = target=20 genes, including Hes2 and, to a lesser degree, Hes4 and Hes5, was=20 increased after radiation exposure (Fig. 1B,=20 and supporting information Figure 1D-1G). Notch signaling in = glioma stem=20 cells was blocked by GSIs,=20 = N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglyc= ine=20 t-butyl ester (DAPT), or L685,458, as shown by = Notch-responsive=20 luciferase reporter assays (Fig. 1C).=20 Using CD133+ glioma cells derived from T4302 and other glioma = sources,=20 including T3359, T4105, and T4597 (Fig. 1D=20 and data not shown), we showed that GSIs moderately reduced growth = of=20 CD133+ glioma cells in the absence of radiation (Fig. 1D,=20 graph 1) but severely impaired growth of cells exposed to = radiation at=20 either 1, 2, or 3 Gy (Fig. 1D,=20 graphs 2-4). On the fifth day after 2 Gy radiation, total cell = numbers in=20 the sham-treated groups were approximately fourfold more than that = in=20 GSI-treated groups. In contrast, without radiation treatment, = sham-treated=20 cells were only 25% (vs. DAPT-treated) or 60% (vs. = L685,458-treated) more=20 than GSI-treated cells 5 days after radiation.=20

    Figure 1. -Secretase inhibitors (GSIs) reduce cell growth = and=20 clonogenic survival of glioma stem cells after radiation.=20 (A): T4302 CD133+ glioma stem cells were transfected = with a=20 Notch-responsive luciferase reporter (RBP-J reporter) premixed with the vector directing = constitutive=20 expression of Renilla luciferase. Cells were left=20 unirradiated or irradiated at 3 Gy and were then collected = to=20 measure luciferase activities at the time points as = indicated. In=20 this and subsequent luciferase assays, firefly luciferase = activity=20 was normalized to the internal control Renilla = luciferase=20 activity, and the normalized firefly luciferase activity = from the=20 sham-irradiated control group was assigned a value of 1. = Data are=20 presented as the mean =B1 standard error. (B): T4302 = CD133+=20 glioma stem cells were left unirradiated or irradiated at 3 = Gy. The=20 relative mRNA levels of Hes2 were determined by quantitative = real-time polymerase chain reaction at the time points as = indicated.=20 (C): T4302 CD133+ cells were transfected with the = RBP-J reporter and treated with dimethyl sulfoxide = (vehicle), 2=20 M DAPT, or 0.5 M L685,458 overnight, and cells = were then=20 collected for luciferase assays. (D): T4302 CD133+ = cells were=20 pretreated with 2 M DAPT or 0.5 M L685,458 for 4 hours and = irradiated as=20 indicated. Total viable cell numbers were then determined = with a=20 cell viability assay for the following 5 days. The averaged = cell=20 titers on day 1 of the control groups were assigned a value = of 1.=20 (E): T4302 CD133+ cells were pretreated and = irradiated as=20 described in (D). Cells were then plated to form=20 neurospheres, and the surviving fractions were determined as = described in Materials=20 and Methods. The mean =B1 standard error was derived = from eight=20 replicates within a representative experiment. (F,G): = Neurosphere formation of T3359 and T4105 CD133+ cells treat = with=20 GSIs alone, 3 Gy radiation alone, or the combination was = determined=20 as described in (E). *P < .05 by Student's = t=20 test; #P < .001 by one-way ANOVA. = Abbreviation:=20 DAPT,=20 = N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglyc= ine=20 t-butyl ester.
    [Normal View 55K | = Magnified View=20 156K]

    Clonogenic survival assay is the standard assay to assess = radiation=20 responsiveness in vitro. In this study, we used neurosphere = formation as a=20 surrogate marker of the clonogenic survival of glioma stem cells. = In=20 combination with radiation, GSIs significantly reduced clonogenic = survival=20 of the T4302 CD133+ glioma cells compared with the sham-treated = groups for=20 all radiation doses examined (Fig. 1E;=20 P =3D .012 at 1 Gy and P < .001 at 2 and 3 Gy by = one-way=20 ANOVA). Similar results were demonstrated using CD133+ cells = derived from=20 other glioma sources, such as T3359, T4105, and T4597 (Fig. 1F,=20 1G,=20 and supporting information Figure 2A, 2B). Taken together, these = results=20 strongly suggest that Notch inhibition with GSIs compromises=20 radioresistance of glioma stem cells.

    GSIs Enhance = Radiation-Induced=20 Cell Death in Glioma Stem Cells

    Significantly greater radiation-induced cell death was observed = in=20 GSI-treated cells compared with sham-treated cells. Two days after = 3 Gy=20 radiation, sham-treated T4302 CD133+ cells contained approximately = 30%=20 dead cells, whereas 60% of cells were dead in the presence of DAPT = or=20 L685,458 (Fig. 2A).=20 Conversely, GSIs did not have an obvious impact on the percentage = of dead=20 cells in the absence of radiation (Fig. 2A).=20 The majority of GSI-treated cells died at 72 hours after 3 Gy of=20 irradiation. However, an excessive amount of cell debris = interfered with=20 the trypan blue exclusion assay at this time point. Activation of = caspase=20 is a hallmark of apoptotic cell death and is critically involved = in=20 radioresponse [[43-46]].=20 Using luminescence-based caspase-3/7 assays, we examined how Notch = inhibition affected radiation-induced caspase activation as a = surrogate=20 marker of cell death. GSIs alone did not significantly alter = caspase=20 activities in the sham-irradiated T4302 CD133+ cells (Fig. 2B;=20 P > .05 by one-way ANOVA). In contrast, combining GSIs = with 3 Gy=20 radiation increased caspase activities by approximately twofold at = 48=20 hours and by fourfold at 72 hours (Fig. 2B;=20 P < .01 by one-way ANOVA). Similar results were detected = using=20 CD133+ glioma cells derived from several other glioma sources, = including=20 T3359, T3691, T4105, and T4597 (supporting information Figure 3 = and data=20 not shown). In validation studies, apoptotic cell death was = assessed by=20 flow cytometry analysis of annexin V, a cell surface marker for = apoptosis.=20 Both DAPT and L685,458 significantly increased annexin V-positive = labeled=20 cells at 48 hours after radiation (Fig. 2C;=20 P < .01 by one-way ANOVA), although the effects were not = obvious=20 at 24 hours after radiation (Fig. 2C;=20 P > .05 by one-way ANOVA). Additionally, DAPT enhanced=20 radiation-induced cell death at different radiation doses (Fig. 2D).=20 The radiosensitizing effects of DAPT showed a = concentration-dependent=20 effect, with 200 nM DAPT (50% inhibition concentration as = suggested by the=20 manufacturer) still being effective (Fig. 2E).=20 Collectively, these data demonstrate that Notch inhibition using = GSIs=20 sensitize glioma stem cells to radiation-induced cell death.=20

    Figure 2. -Secretase inhibitors (GSIs) enhance = radiation-induced cell=20 death in glioma stem cells. (A): T4302 CD133+ cells = were=20 pretreated with DAPT or L685,458 for 4 hours and irradiated = at 3 Gy.=20 Percentage of dead cells was determined with trypan blue = exclusion=20 assay at 48 hours after radiation exposure. (B): = T4302 CD133+=20 cells were treated with GSIs =B1 radiation as indicated. = Relative=20 caspase 3/7 activities were determined by normalizing = caspase=20 activities to the corresponding cell titers. (C): = T4302=20 CD133+ cells were treated as described in (A).At 24 = or 48=20 hours after radiation exposure, cells were labeled with=20 FITC-conjugated annexin V staining and analyzed with flow = cytometry.=20 (D): T4302 CD133+ cells were pretreated with 2 M DAPT =B1 radiation as = indicated. Relative=20 caspase 3/7 activities were determined at 72 hours after = radiation.=20 (E) T4302 CD133+ cells were treated with DAPT at = indicated=20 concentrations for 4 hours and irradiated at 3 Gy. Relative = caspase=20 activities were determined at 48 or 72 hours after = radiation.=20 *P < .01 by Student's t test; = #P=20 < .01 by one-way ANOVA. Abbreviation: DAPT,=20 = N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglyc= ine=20 t-butyl ester.
    [Normal View 48K | = Magnified View=20 128K]

    Expression of the=20 Constitutively Active Intracellular Domains of Notch1 or Notch2 = Attenuates=20 the Radiosensitizing Effects of GSIs

    Substrates of GSIs include more than 50 type I membrane = proteins in=20 addition to the Notch receptors [[47]].=20 The radiosensitizing effects of GSIs are not necessarily mediated = through=20 inhibition of Notch. We therefore validated the role of Notch in=20 radioresponse of glioma stem cells by expression of the = constitutively=20 active intracellular domains of Notch1 or Notch2 (NICD1 or NICD2) = that=20 function downstream of GSIs. Activation of the Notch pathway by=20 lentivirus-mediated expression of NICD1 or NICD2 was validated by=20 upregulation of Notch target genes, including Hes2, Hes4, and Hes5 = (Fig.=20 3A),=20 as well as increased luciferase activities generated from the=20 Notch-responsive reporter (Fig. 3B).=20 Similar to the parental cells, cells infected with the control = lentivirus=20 were sensitive to DAPT when irradiated (Fig. 3C).=20 In contrast, cells expressing either NICD1 or NICD2 demonstrated=20 significantly weaker radiation-induced cell death in comparison to = control=20 cells regardless of DAPT treatment (Fig. 3C).=20 Additionally, DAPT treatment failed to increase the percentage of = annexin=20 V-positive cells in the presence of NICD2 expression (Fig. 3D).=20 Finally, expression of NICD2 improved clonogenic survival of the = T4302=20 glioma stem cells after radiation irrespective of DAPT treatment = (Fig. 4E;=20 P =3D .004, NICD2 vs. control =B1 DAPT; P < .01, = NICD2 + DAPT=20 vs. control =B1 DAPT, at 3 Gy radiation by one-way ANOVA), whereas = DAPT=20 treatment did not significantly alter clonogenic survival of = irradiated=20 NICD2-expressing cells (Fig. 4E;=20 P > .05, NICD2 vs. NICD2+DAPT, by Student's t = test).=20 Additionally, expression of NICD1 or NICD2 protected glioma stem = cells=20 derived from T3359 and T4105 in a similar manner (supporting = information=20 Figure S4 and data not shown). Taken together, our data = demonstrate that=20 Notch activation by expression of NICDs attenuates the = radiosensitizing=20 effects of GSIs, suggesting that these activities of GSIs are = crucially=20 mediated through inhibition of Notch.=20

    Figure 3. Expression = of the=20 intracellular domains of Notch1 or Notch2 attenuates the=20 radiosensitizing effects of -secreting inhibitors (GSIs). T4302 CD133+ glioma = cells=20 were infected with control lentivirus or lentivirus = directing=20 expression of NICD1 or NICD2 and selected by puromycin. = (A):=20 Expression of Notch1, Notch2, Hes2, Hes4, and Hes5 were = determined=20 by quantitative real-time polymerase chain reaction. = (B):=20 Control cells or cells expressing NICD2 were transfected = with=20 Notch-dependent luciferase reporter and treated with = dimethyl=20 sulfoxide or 2 M DAPT overnight. = Notch-dependent=20 transcription activities were determined by luciferase = reporter=20 assays as described in Figure 1.=20 (C): Control cells or cells expressing NICDs were = treated=20 with DAPT =B1 radiation as indicated. Relative caspase 3/7 = activities=20 were determined at 48 or 72 hours after radiation. = (D):=20 Apoptotic cell death was determined with annexin V staining = as=20 described in Figure 2.=20 (E): Surviving fractions were determined as described = in=20 Figure 1.=20 *P < .01 by Student's t test; = #P=20 < .001 by one-way ANOVA. Abbreviations: DAPT,=20 = N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglyc= ine=20 t-butyl ester; NICD, Notch receptor intracellular=20 domain.
    [Normal = View=20 46K | Magnified = View=20 125K]

    Figure 4. Notch = pathway does not=20 alter DNA damage response of glioma stem cells. (A): = T4302=20 CD133+ cells were pretreated with DAPT or L685,458 for 24 = hours=20 before radiation. At 1 hour after radiation exposure, cells = were=20 harvested. The levels of activating phosphorylation of Chk1 = and Chk2=20 were assessed by immunoblotting. (B): T4302 CD133+ = cells were=20 treated with 2 M DAPT at 4 and 24 hours before = radiation=20 (-24 hours or -4 hours) or at 0, 4, and 24 hours after = radiation.=20 Relative caspase 3/7 activities were measured 3 days after = radiation=20 exposure. *P < .001 by Student's t test.=20 Abbreviation: DAPT,=20 = N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglyc= ine=20 t-butyl ester.
    [Normal View 25K | = Magnified View=20 66K]

    Notch Pathway Does = Not Alter=20 DNA Damage Response of Glioma Stem Cells

    The DNA damage checkpoint response plays a critical role in = cellular=20 response to radiation [[48][49]].=20 Our previous study showed that increased activation of the DNA = damage=20 response is implicated in radioresistance of the glioma stem cells = [[7]].=20 To determine the mechanisms through which Notch promotes = radioresistance=20 of glioma stem cells, we first assessed whether the Notch pathway = affected=20 activation of the checkpoint kinases after radiation. Neither DAPT = nor=20 L685,458 significantly altered activating phosphorylation of Chk1 = or Chk2=20 after radiation (Fig. 4A).=20 Additionally, in either T4105 or T4302 CD133+ glioma cells, DAPT = increased=20 radiation-induced cell death even when it was added 24 hours after = radiation (Fig. 4B,=20 and supporting information Figure 5), whereas the bulk DNA damage = repair=20 process is usually completed within a few hours after radiation = exposure=20 [[50]].=20 These results suggest that the DNA damage response pathway is not=20 critically involved in the radioprotective activities of = Notch.

    Notch Promotes = Radioresistance=20 Through Regulation of the PI3K/Akt Pathway and Mcl-1

    Notch inhibition renders glioma stem cells more sensitive to=20 radiation-induced cell death. In a context-dependent manner, Notch = can=20 promote cellular survival through different mechanisms, such as = activation=20 of the PI3K/Akt pathway [[51][52]]=20 and upregulation of the prosurvival proteins Bcl-2 and Mcl-1 [[53]].=20 We compared alterations of these prosurvival factors between the=20 NICD2-expressing glioma stem cells and the control cells treated = with=20 DAPT, radiation, or both. Activating phosphorylation at serine 473 = of Akt=20 was used as a surrogate marker of Akt activity. While radiation = exposure=20 induced Akt activation as shown by increased levels of = phospho-S473 Akt,=20 DAPT treatment reduced Akt activation (Fig. 5A).=20 In contrast, expression of NICD2 increased Akt activities, which = was=20 partially attenuated but not abolished by DAPT (Fig. 5A).=20 Similar results were found in other short-term glioma cultures, = such as=20 T3359 and T4105 (data not shown). Our laboratory recently reported = that=20 Akt activity was essential for survival of glioma stem cells [[37]].=20 In agreement with this finding, treatment of the PI3K inhibitor = LY294002=20 or the Akt inhibitor III increased glioma stem cell death with or = without=20 radiation (Fig. 5B).=20 Treatment with LY294002 or the Akt inhibitor reduced = phosphorylation at=20 S473 of Akt (supporting information Figure 6). Expression of NICD2 = did not=20 overcome the apoptotic effects of either LY294002 or the Akt = inhibitor=20 III, suggesting that the PI3K/Akt pathway functions downstream of = Notch=20 (Fig. 5B).=20

    Figure 5. Notch = pathway=20 regulates radioresistance of glioma stem cells via the Akt = pathway=20 and Mcl-1. T4302 CD133+ cells were infected with control = lentivirus=20 or lentivirus directing expression of NICD2. Cells were = pretreated=20 with or without 2 M DAPT for 4 hours and exposed = to 3 Gy=20 radiation. Irradiated cells were collected at 24 or 48 hours = after=20 radiation, and sham-irradiated cells were collected at 48 = hours=20 after radiation. (A, C, D): Protein levels of = phospho-Akt=20 (S473), total Akt, PTEN, Bcl-2, Bcl-xL, Mcl-1, phospho-p53 = (S15),=20 total p53, and PUMA were assessed by immunoblotting. Actin = was=20 blotted as the loading control. (B): Control cells or = NICD2-expressing cells were pretreated with 10 M PI3K inhibitor, 10 M LY294002, or 50 M Akt inhibitor III (SH-6), for = 4 hours=20 before radiation exposure. Relative caspase 3/7 activities = were=20 determined at 48 hours after radiation exposure.=20 #P < .001 by one-way ANOVA. = Abbreviations:=20 DAPT,=20 = N-[N-(3,5-difluorophenacetyl)-l-alanyl]-S-phenylglyc= ine=20 t-butyl ester; NICD, Notch receptor intracellular=20 domain.
    [Normal = View=20 50K | Magnified = View=20 160K]

    The anti-apoptotic Bcl-2 proteins, such as Bcl-2, Bcl-xL, and = Mcl-1,=20 are important promoters for cell survival [[54]].=20 The levels of Bcl-2 and Bcl-xL were not apparently altered by = either=20 radiation or Notch alteration (Fig. 5C).=20 Conversely, radiation exposure or DAPT treatment decreased the = levels of=20 Mcl-1, and expression of NICD2 increased Mcl-1 levels even in the = presence=20 of DAPT (Fig. 5C).=20 Interestingly, radiation or DAPT increased the levels of Mcl-1 = proteins of=20 a smaller size (Fig. 5C).=20 These bands could represent either proteolytically cleaved Mcl-1 = proteins=20 generated during apoptosis or a truncated isoform of Mcl-1=20 (Mcl-1S), which is pro-apoptotic due to loss of BH1 and = BH2=20 domains [[55]].

    The functional interaction between Notch and p53 is intriguing, = as=20 Notch can activate p53 [[56][57]]=20 or inhibit p53 [[58]]=20 in a cell type-specific manner. As a crucial regulator of cellular = radioresponse, p53 activity may be involved in the radioprotective = function of Notch. We saw that expression of NICD2 increased p53 = levels in=20 the absence of radiation and further enhanced p53 activation = induced by=20 radiation (Fig. 5D).=20 Additionally, expression of the p53 target gene PUMA was = upregulated by=20 NICD2 (Fig. 5D).=20 These results are consistent with a previous study showing that = Notch1=20 upregulates p53 in gliomas [[57]]=20 and suggest that the radioprotective functions of Notch are not = mediated=20 by inhibition of p53.

    CD133-Negative = Glioma Cells=20 Do Not Respond to GSIs

    We further examined whether Notch inhibition affected = radioresponse of=20 the CD133-negative fraction of gliomas. After similar procedures, = cell=20 death and clonogenic survival after radiation exposure were = determined in=20 the T3359 and T4302 CD133-negative cells with and without GSI = treatment.=20 In contrast to glioma stem cells, caspase activation or clonogenic = survival of the CD133-negative glioma cells were not significantly = altered=20 by Notch inhibition using GSIs irrespective of radiation exposure = (Fig. 6A-6D).=20 Additionally, GSI treatment did not alter Akt activities or Mcl-1 = levels=20 in the CD133-negative glioma cells (Fig. 6E).=20 Taken together, our data suggest that the radiosensitizing effects = of=20 Notch inhibition are specific to the cancer stem cell fraction of = gliomas.=20

    Figure 6. Notch = inhibition by=20 -secreting inhibitors (GSIs) does not = significantly alter=20 radiation response of CD133-negative glioma cells. T3359=20 CD133-negative cells were pretreated with 2 M GSI or 0.5 M L685,458 for 4 hours and = irradiated at 3=20 Gy or as indicated. (A): Relative caspase 3/7 = activities were=20 determined at 48 or 72 hour after radiation as described in = Figure=20 2.=20 (B): After radiation, cells in 6-well plates were = cultured=20 for 3 weeks and then fixed and stained to count colonies. = Surviving=20 fractions were determined as described in Materials=20 and Methods. (C, D): The effects of GSIs on cell = death=20 and clonogenic survival of T4302 CD133-negative cells were=20 determined as described above. ##P > = .05 by=20 one-way ANOVA. (E): T4302 CD133-negative cells were = treated=20 with GSIs =B1 radiation as described above. Levels of = phospho-Akt=20 (S473), total Akt, Mcl-1, and actin were determined by=20 immunoblotting.
    [Normal=20 View 49K | Magnified View=20 139K]

    Knockdown of = Notch1 or Notch2=20 Sensitizes Glioma Stem Cells to Radiation and Impairs Xenograft = Tumor=20 Formation

    Individual Notch receptors may have distinct biological = functions in=20 certain tumor types [[59]].=20 It has been demonstrated that expression of the constitutively=20 intracellular regions of Notch1 or Notch2 appears to exert = opposite=20 effects in medulloblastomas, with Notch1 acting as a tumor = suppressor and=20 Notch2 acting as a tumor promoter [[60]].=20 In gliomas, knockdown of Notch1 impairs cell growth and survival = [[61]].=20 By expression of NICD1 or NICD2, our results suggest that both = Notch=20 receptors promote radioresistance of glioma stem cells. To gain a = better=20 understanding of how individual Notch receptors are involved in=20 radioresponse, we determined the effects of knockdown of Notch1 or = Notch2=20 on radioresistance through lentivirus-mediated expression of shRNA = sequences targeting Notch1 or Notch2. The efficiency of = downregulation of=20 the corresponding Notch receptors and Notch target genes were = determined=20 by quantitative real-time polymerase chain reaction (Fig. 7A).=20 Consistent with previous report, knockdown of Notch1 alone = increased=20 apoptotic cell death in T3359 and T4302 CD133+ cells. = Interestingly,=20 similar effects were achieved by knockdown of Notch2 (Fig. 7B,=20 and supporting information Figure 7A). Cells with reduced Notch1 = or Notch2=20 expression were highly sensitive to radiation-induced cell death = (Fig. 7B,=20 and supporting information Figure 7A). Consistently, the = postradiation=20 clonogenic survival of glioma stem cells was severely impaired by=20 knockdown of either Notch1 or Notch2 (Fig. 7C,=20 and supporting information Figure 7B). When exposed to 3 Gy = radiation,=20 glioma stem cells expressing shRNA specific to Notch1 or Notch2 = rarely=20 formed neurospheres (Fig. 7C,=20 7D).=20 Additionally, knockdown of Notch1 or Notch2 alone reduced cell = viability=20 of the CD133-negative glioma cells but did not significantly = increase=20 radiation-induced cell death (supporting information Figure 7C; = P=20 > .05, irradiated groups vs. corresponding sham-irradiated = groups by=20 Student's t test). These results further validate the = specific role=20 of Notch pathway in regulation of radioresistance and suggest that = multiple Notch receptors may be involved in radiation response of = glioma=20 stem cells.=20

    Figure 7. Knockdown of = Notch1 or=20 Notch2 increases radiosensitivity of glioma stem cells in = vitro and=20 in vivo. T4302 CD133+ cells were infected with lentivirus = directing=20 expression of nontargeting shRNA or shRNAs specific to = Notch1 or=20 Notch2 (KD-Notch1 or KD-Notch2) and selected with puromycin. = (A): Relative mRNA levels of Notch1, Notch2, Hes4, or = Hes5=20 were determined with quantitative real-time polymerase chain = reaction. *P < .05 by Student's t test. = (B):=20 Relative caspase 3/7 activities were determined in control = cells or=20 cells with Notch knockdown treated with or without = radiation.=20 #P < .001 by one-way ANOVA. (C): = Clonogenic survival was compared between the control group = and cells=20 with Notch knockdown as described in Figure 1.=20 (D): Representative images (=D7100) of neurospheres = formed by=20 T4302 CD133+ cells with or without Notch knockdown 2 weeks = after=20 radiation. (E): After lentivirus infection and = puromycin=20 selection, T4302 CD133+ cells were left unirradiated or = irradiated=20 at 3 Gy, and they were immediately injected into both flanks = of=20 athymic nude mice at 250,000 cells per injection. Three mice = were=20 inoculated for a total of six injections per group. Tumor = growth was=20 monitored twice a week for 4 weeks until diameters of the = largest=20 tumors reached approximately 20 mm.
    [Normal View 57K | = Magnified View=20 178K]

    We examined how impaired Notch signaling affected radiation = response of=20 glioma stem cells in vivo using a flank xenograft model. Control = T4302=20 CD133+ glioma cells or cells with reduced Notch1 or Notch2 = expression were=20 subcutaneously inoculated into both flanks of nude mice without = radiation=20 exposure or immediately after 3 Gy radiation exposure. The = sham-irradiated=20 control cells formed palpable tumors at all injection sites at day = 14=20 after injection (Fig. 7E,=20 and supporting information Table 1). Knockdown of either Notch1 or = Notch2=20 alone modestly extended tumor latency (Fig. 7E).=20 Tumors formed by irradiated control cells were first detected at = day 17,=20 and all became palpable at day 21 (Fig. 7E).=20 In combination with radiation, knockdown of Notch1 or Notch2 not = only=20 extended the tumor latency (Fig. 7E)=20 but also significantly decreased tumor incidence (1 of 6 for = knockdown of=20 Notch1, and 2 of 6 for knockdown of Notch2; supporting information = Table=20 1). Taken together, our results suggest that integrated Notch = signaling is=20 critically implicated in radiation resistance of glioma stem cells = in=20 vivo.

    DISCUSSION
    3DDISCUSSION=20

    Fundamental improvements in brain cancer treatment will require = the=20 development of new therapeutic paradigms. Temozolomide, an oral=20 methylating chemotherapeutic agent, became the standard of care = for newly=20 diagnosed glioblastomas when used concurrently with external beam=20 radiation followed by adjuvant therapy [[3]],=20 suggesting that combining chemotherapies with radiation may shift = the=20 survival curve. Despite the benefit of temozolomide, glioblastomas = continue to be highly resistant to radiation [[4]].=20 We recently demonstrated that the cancer stem cell fraction of = gliomas was=20 more resistant to radiation than non-stem glioma cells [[7]].=20 Given the high tumorigenic capacity of glioma stem cells, this = paradigm=20 suggests that disrupting radioresistance of glioma stem cells may = augment=20 the efficacy of radiotherapy. We have already shown that = inhibitors of the=20 checkpoint kinases, Chk1 and Chk2, sensitized glioma stem cells to = radiation [[7]].=20 However, DNA damage checkpoint inhibitors may have limited = therapeutic=20 index due to shared dependence of normal cells on these molecules = in=20 radioresponse [[62]].=20 In this study, we described a novel radioprotective role of Notch=20 signaling in glioma stem cells. Our results show that radiation = induces=20 Notch activation, and Notch inhibition by GSIs or = Notch1/2-specific shRNA=20 renders glioma stem cells more sensitive to radiation. In further = support=20 of these observations, activation of Notch via expression of NICD1 = or=20 NICD2 promotes radioresistance of glioma stem cells. Collectively, = these=20 pharmacological and genetic approaches show that Notch activity is = critically implicated in radioresistance of glioma stem cells, = suggesting=20 that the Notch pathway may serve as a potential therapeutic target = for=20 improvement of radiotherapy against gliomas.

    The oncogenic role of Notch is highlighted by the presence of = mutations=20 of the Notch pathway components in a variety of human tumors, in=20 particular leukemia and breast cancer [[63]].=20 Upregulation of Notch pathway components also presents in = glioblastoma=20 multiforme cell lines and surgical biopsy specimens [[30]].=20 A critical role of the Notch pathway in the maintenance of brain = tumor=20 stem cells has recently emerged [[41]].=20 Our results suggest that the radioprotective functions of Notch = are=20 specific to the CD133+ glioma stem cells. Neither cell death nor=20 clonogenic survival of the non-stem glioma cells are significantly = altered=20 by GSI treatment irrespective of radiation exposure. Knockdown of = Notch1=20 or Notch2 modestly impaired cell viability in the non-stem glioma = cells,=20 but did not alter radiation-induced cell death. These studies = suggest that=20 stem cell-specific pathways, such as Notch, can serve as potential = therapeutic targets that may not be readily appreciated by studies = using=20 whole tumors.

    The Notch pathway can promote or repress tumorigenesis in a=20 context-dependent manner. For example, loss of Notch1 or = expression of a=20 dominant negative MAML1 mutant in mouse epidermis induces = hyperplasia and=20 subsequent skin tumors [[64][65]].=20 In medulloblastomas, expression of NICD2 promotes tumor growth, = whereas=20 NICD1 is tumor suppressive [[60]].=20 Conversely, knockdown of Notch1 impaired proliferation and = survival of=20 glioma cell lines [[61]].=20 Using lentivirus-mediated expression of NICD1/2 or multiple=20 Notch1/2-specific shRNAs, our data suggest that both Notch1 and = Notch2=20 promote radioresistance in glioma stem cells, although the = relative=20 contribution of each Notch receptor is undefined. It remains to be = elucidated why knockdown of individual Notch receptors leads to = moderate=20 loss of cell viability in glioma stem cells in the absence of = radiation,=20 whereas blockage of Notch signaling using GSIs only reduces cell = growth.=20 These effects may represent sustained inhibition of Notch by = stable=20 expression of shRNAs, off-target activities of the shRNAs, or=20 oversaturation of the microRNA/shRNA pathways [[66]].=20 Alternatively, membrane-bound Notch receptors may have important=20 biological functions in gliomas that have not been fully = recognized, as a=20 recent report suggests that membrane-anchored Notch1 activates Akt = in a=20 transcription-independent manner [[67]].=20 The specific role of each individual Notch receptor in regulation = of=20 radioresistance of glioma stem cells is beyond the scope of this = study,=20 but there is substantial interest with regard to the therapeutic = potential=20 of Notch inhibition because specific Notch neutralizing antibodies = are=20 under clinical development for systemic cancer treatment and may = have=20 distinct adverse effects in a therapeutic setting. Notch1 is a key = regulator of intestinal stem cell fate determination and gut = homeostasis=20 [[22][23]].=20 Clinical trials using GSIs for treatment of Alzheimer's disease = are=20 primarily associated with gastrointestinal toxicity [[68][69]],=20 which is likely due to inhibition of Notch1 by GSIs. If Notch = receptors=20 other than Notch1 are also essentially required for = radioresistance of=20 glioma stem cells, small molecules or antibodies that specifically = target=20 these receptors while sparing Notch1 may achieve similar = radiosensitizing=20 effects in gliomas with reduced gut toxicity.

    Our results demonstrate that Notch inhibition in irradiated = glioma stem=20 cells dramatically increased cell death within 3 days after = radiation,=20 suggesting that Notch is crucially involved in postradiation = survival of=20 glioma stem cells. In accordance with these observations, Notch=20 upregulates the prosurvival factors Akt and Mcl-1 in glioma stem = cells. It=20 has been reported that Mcl-1 is a critical regulator of survival = in neural=20 precursor cells during development and in response to DNA damage = [[53][70]].=20 Akt is known to upregulate Mcl-1 levels [[71][72]].=20 Therefore, the regulation of Mcl-1 by Notch can be potentially = mediated=20 through Akt. The PI3K/Akt pathway plays a central role in the=20 radioprotective functions of Notch, as shown by our results that=20 inhibitors of PI3K or Akt abolish the radioprotective activities = of NICD2.=20 This pathway has profound functions in many aspects of cell = growth,=20 proliferation, and survival [[73]].=20 We previously described an important link between the PI3K/Akt = pathway=20 with growth and survival of glioma stem cells [[37]].=20 Confirmation of these findings came from studies of a mouse=20 medulloblastoma model in which the PI3K/Akt pathway is activated = in=20 medulloblastoma stem cells after radiation and is essentially = required for=20 postradiation cell survival [[74]].=20 In addition, inhibition of Akt sensitizes the medulloblastoma stem = cells=20 to radiation [[74]].=20 The mechanisms through which Notch activates the PI3K/Akt pathway = in=20 glioma stem cells remain to be elucidated. Although Notch can = activate Akt=20 through negatively regulated PTEN in T-cell acute lymphoblastic = leukemia=20 [[51]],=20 our results show that expression of NICD2 upregulated PTEN = irrespective of=20 radiation exposure, suggesting that Notch does not activate the = PI3K/Akt=20 pathway by repressing PTEN. Alternatively, the Src-related = tyrosine kinase=20 p56lck has been reported to mediate Notch-induced Akt activation = [[52]].=20 Our results show that neither a p56lck-specific inhibitor not a = pan-Src=20 family kinase inhibitor altered radiation response of glioma stem = cells=20 (data not shown), suggesting that Src family tyrosine kinases are = not=20 implicated in radiation response of glioma stem cells. = Interestingly, a=20 recent study reports a non-canonical transcription-independent = Notch=20 function that membrane-anchored Notch1 is sufficient to activate = Akt [[67]].=20 This novel mechanism may be implicated in Akt activation by Notch = in=20 glioma stem cells, but details of this functional interaction = remain to be=20 determined. Our data show that NICD2 upregulated PTEN and p53 in = glioma=20 stem cells instead of repressing these two tumor suppressors, = suggesting=20 that these two proteins may not be directly involved in the=20 radioprotective functions of Notch. However, postradiation = survival of=20 medulloblastoma stem cells can be promoted by cell cycle arrest = mediated=20 through PTEN and p53 [[74]].=20 Therefore, it is possible that, in glioma stem cells, = Notch-induced PTEN=20 and p53 indirectly contribute to reduced cell death after = radiation by=20 promoting cell cycle arrest. Taken together, our study suggests a=20 Notch-regulated signaling pathway that is critically involved in = radiation=20 response of glioma stem cells. This pathway provides multiple = potential=20 therapeutic targets at several levels, including the Notch = receptors, the=20 kinases (PI3K, Akt), and Mcl-1, a molecule that directly regulates = the=20 apoptosis cascade.

    The last proteolytic step that generates the active = intracellular=20 domains of Notch receptors is dependent on the -secretase complex [[18][19]].=20 A substantial number of GSIs have been developed for treatment of=20 Alzheimer's disease because the -secretase complex mediates production of -amyloid peptides, the precursor of = amyloid=20 plaques found in Alzheimer's disease [[75]].=20 Given their activities of inhibiting Notch, GSIs have also been = actively=20 examined for their anti-cancer efficacy in a variety of tumor = types, such=20 as T-cell acute lymphoblastic leukemia [[76]],=20 breast cancer [[77]],=20 Kaposi's sarcoma [[78]],=20 medulloblastoma [[79]],=20 intestinal adenoma [[23]],=20 etc. Clinical trials with the GSI MK0752 (Merck & Co., = Whitehouse=20 Station, NY, http://www.merck.com/) are underway in patients = with=20 breast cancer and pediatric central nervous system malignancies = (http://clinicaltrials.gov/). Our study suggest = that GSIs=20 alone only display limited growth inhibitory effects in glioma = stem cells;=20 in combination with radiotherapy, however, GSIs may provide = significant=20 therapeutic benefits by promoting the radiosensitivity of glioma = stem=20 cells. Therefore, additional efforts will be required to evaluate = the=20 clinical benefits of GSIs in combination with radiation as the = monotherapy=20 trials of GSIs may not show significant antitumor efficacy in = tumors such=20 as gliomas. Taking account into the prevalent role of Notch in = regulation=20 of stem cell behavior and cancer biology, it is reasonable to = speculate=20 that the radiosensitizing effects of Notch inhibition are not = limited to=20 gliomas and may be a broadly useful treatment paradigm.

    Disclosures of Potential Financial = Conflicts=20 of Interest

    The authors indicate no potential conflicts of interest.

    Acknowledgements
    3DAcknowledgements=20

    The authors thank Spyros Artavanis-Tsakonas, Charles Eberhart, = and=20 Adolfo Ferrando for reagents. We are also grateful to Mark = Dewhirst for=20 helpful discussion. Jialiang Wang is a recipient of the Basic = Research=20 Fellowship from the American Brain Tumor Association. Rebekah = White is a=20 recipient of the Howard Hughes Medical Institute Early Career = Award.=20 Justin Lathia is a recipient of a Basic Research Fellowship from = the=20 American Brain Tumor Association and a National Service Research = Award=20 (NCI F32 CA142159). Support for Bruce Sullenger came from National = Institutes of Health (NIH) grant CA129190, and support for Jeremy = Rich=20 came from the Childhood Brain Tumor Foundation, the Pediatric = Brain Tumor=20 Foundation of the United States, Accelerate Brain Cancer Cure, = Alexander=20 and Margaret Stewart Trust, Brain Tumor Society, Goldhirsh = Foundation,=20 Sidney Kimmel Foundation, Damon Runyon Cancer Research Foundation, = and NIH=20 grants NS047409, NS054276, CA129958, CA116659. The funders had no = role in=20 study design, data collection and analysis, decision to publish, = or=20 preparation of the manuscript.

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