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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_0010_01CA2894.856E2720 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://download.interscience.wiley.com/freeflow/js/personalisation.js /*=0A= * Project : Interscience Redesign=0A= * File : personalisation.js=0A= * Creation Date : 25th March 2003=0A= * Author : Jonathan Sugar=0A= *=0A= * Various functions to deliver personalisation functionality.=0A= */=0A= =0A= // Pre-declare userName so it's available when we need to test if the = page=0A= // assembler ran OK=0A= var userName =3D null;=0A= var shcrtCookie =3D null;=0A= =0A= // Browser detect globals:=0A= var uaStr =3D navigator.userAgent.toLowerCase();=0A= var uaVer =3D parseInt(navigator.appVersion);=0A= // for detecting netscape 4=0A= var netscape =3D (uaStr.indexOf("mozilla") > -1);=0A= var netscape4 =3D (netscape && uaVer < 5);=0A= =0A= // Factory Methods=0A= =0A= // Simple wraper object to encapsulate the data for a co-branding item.=0A= function coBrandingItem(image, imageAltText, href)=0A= {=0A= //alert("Making new coBrandingItem:\nImage: " + image + = "\nimageAltText" + imageAltText + "\nhref: " + href);=0A= this.image =3D image;=0A= this.imageAltText =3D imageAltText;=0A= this.href =3D href;=0A= }=0A= =0A= /*=0A= * Insert the 'Customer co-branding' section.=0A= * -----------------------------=0A= *=0A= * Author: Jonathan Sugar, March 2003.=0A= *=0A= * 1. Insert customer co branding. Loop through 'customerCoBranding' array=0A= * inserting XHTML where needed.=0A= *=0A= * NOTE: the array contains ONLY the resource location (the location of a = gif=0A= * on the file system), the required XHTML is then added where needed. = This way=0A= * we maintain the seperation between presentation and content.=0A= */=0A= function insertCustomerCoBranding(maxBranding)=0A= {=0A= if( customerCoBranding[0].image !=3D "none")=0A= {=0A= document.writeln('

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

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

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

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

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

3D""

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

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

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

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

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

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

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

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

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

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

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

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

";=0A= }=0A= =0A= } ------=_NextPart_000_0010_01CA2894.856E2720 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://download.interscience.wiley.com/freeflow/js/site.js /************************************************************************= ***************************=0A= FILE: site.js=0A= AUTHOR: Jonathan Sugar, Chris Neatby-Smith and Digital Pulp. July = 2003.=0A= =0A= DESCRIPTION: Javascript relating to site wide functionality across =0A= InterScience.=0A= =0A= INCLUDES FUNCTIONS=0A= =3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=0A= =0A= Search Functions=0A= ----------------=0A= =0A= selectAndSubmit() - for deciding where to submit a query based on the = state of the form.=0A= processallContentSearch() - for validating form input for an all = content search.=0A= processTitleSearch() - for validating form input for a Title search.=0A= checkCitationSearch() - for validating the citation search form.=0A= selectAll() - check all the checkboxes in a form.=0A= do_popup() - stasa Acronym Finder window popup function.=0A= =0A= =0A= Other Functions=0A= ---------------=0A= =0A= fixInlineLists() - fix netscape 4.x bug where

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

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

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

You do not have = anything saved in this menu.

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

" = + liels[j].innerHTML + "

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

" + liels[j].innerHTML + "

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

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FEBS Journal

ISSN: 1742-4658 (Online)

ISSN: 1742-464X (Print)

Volume 276, Issue 17, 2009.

Pages: 4653=964664

Top of Page
Abstract
Introduction
Angiogenesis in brain tumors
Characteristics of brain tumor vasculature
Targeting brain tumor vasculature
Addendum
Acknowledgements
References

Tables: 1

List of=20 Images

Abstract=20 Only

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3D"About

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

Copyright =C2=A9 2007 Pathological Society of Great = Britain and Ireland. Published by John Wiley & Sons, Ltd.

'); } =0A= function copyright(hwnd) {var title =3D pagetitle(); = hwnd.document.write(title.bold() + '
Copyright Journal compilation = © 2009 Federation of European Biochemical Societies

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'); } =0A= =0A= =0A= function caption_f1(hwnd) = {hwnd.document.write('Fig. 1. Angiogenic interaction = between brain tumor and endothelial cells and antiangiogenic therapy. = VEGF expression is induced in tumor cells by hypoxia [19], radiation, = receptor tyrosine kinase signaling and Akt pathways [114]. VEGF and = other proangiogenic factors are secreted by brain cancer cells in a = paracrine and endocrine manner [83]. These factors support endothelial = cell survival. Also, VEGF induces permeability of the BBB [78]. Notch = signaling mediated by γ-secretase suppresses angiogenesis but is = required for proper vascular development [38–40,44]. The = signaling required for angiogenesis in brain tumors can be targeted = using an antibody against VEGF [20,68], a decoy receptor to VEGF [115], = small-molecule TKIs of VEGFR[82], inhibitors of other kinases in the = signaling cascade and inhibitors of γ-secretase [85]. Ab, = antibody; HGF/SF, hepatocyte growth factor/scatter factor; IR, ionizing = radiation; RTK, receptor tyrosine kinase; PTEN, phosphatase and tensin = homologue; PI3K, phosphatidylinositol-3 kinase; HRE, hypoxia-responsive = element; ROS, reactive oxygen species; γ-Sec, = γ-secretase.'); }=0A= function display_f1(size) { var i =3D new Array(); i[0] =3D 'nf1'; i[1] = =3D 'nf1'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f1', size); imagelink(hwnd, i[size]); = caption_f1(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= =0A= =0A= function imagelist() {var hwnd =3D newfigwin(); dochead(hwnd, 'List of = Images - '); copyright(hwnd); =0A= hwnd.document.write('');=0A= =0A= =0A= hwnd.document.write('');hwnd.document.write('

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Volume 276, Issue 17, Pages 4653-4664

Published Online: 31 Jul 2009

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3D""=20

MINIREVIEW

Brain angiogenesis in developmental = and=20 pathological processes: mechanism and therapeutic intervention in = brain=20 tumors

Woo-Young Kim and=20 Ho-Young Lee

Department of = Thoracic/Head and=20 Neck Medical Oncology, The University of Texas M. D. Anderson = Cancer=20 Center, Houston, TX, USA

Correspondence to H.-Y. Lee, = Department=20 of Thoracic/Head and Neck Medical Oncology, Unit 432, The = University of=20 Texas M. D. Anderson Cancer Center, 1515 Holcombe Boulevard, = Houston, TX=20 77030, USA
Fax: +1 713 792 0430
Tel:=20 +1 713 745 0769
E-mail: hlee@mdanderson.org

Copyright Journal compilation =A9 2009 = Federation=20 of European Biochemical Societies

KEYWORDS

antiangiogenic therapy =95 avarstin =95 BBB = =95 brain tumor=20 angiogenesis =95 clinical trial =95 GBM =95 resistance =95 VEGF = =95 vessel=20 normalization

ABSTRACT

3DAbstract=20

Formation of new blood vessels is required for the = growth=20 and metastasis of all solid tumors. New blood vessels are = established in=20 tumors mainly through angiogenesis. Brain tumors in particular are = highly=20 angiogenic. Therefore, interventions designed to prevent = angiogenesis may=20 be effective at controlling brain tumors. Indeed, many recent = findings=20 from preclinical and clinical studies of antiangiogenic therapy = for brain=20 tumors have shown that it is a promising approach to managing this = deadly=20 disease, especially when combined with other cytotoxic treatments. = In this=20 minireview, we summarize the basic characteristics of brain tumor=20 angiogenesis and the role of known angiogenic factors in = regulating this=20 angiogenesis, which may be targets of antiangiogenic therapy. We = also=20 discuss the current status of antiangiogenic therapy for brain = tumors, the=20 suggested mechanisms of this therapy and the limitations of this=20 strategy.

(Received 19 February 2009, revised 14 May 2009, = accepted 23=20 June 2009)

DIGITAL OBJECT IDENTIFIER (DOI)

10.1111/j.1742-4658.2009.07177.x About=20 DOI

=20

=20

=20

Introduction

Brain tumors are much rarer than tumors of other = organs.=20 Specifically, fewer than 2% of all tumors diagnosed each year are = primary=20 brain tumors [1].=20 Furthermore, because of the location and other physiological=20 characteristics of brain tumors, their prognosis is very poor [1].=20 Whereas primary brain tumors originate in the brain, secondary = brain=20 tumors metastasize from other cancers, such as breast, lung and = colon=20 cancer, and melanoma. Of the many different types of brain tumors, = glioblastoma multiforme (GBM) is the most common, accounting for=20 ∼ 40% of all primary brain tumors and 70% of all = malignant gliomas=20 [2=964].=20 GBM is also one of the most vascular and deadly cancers, with a = very low=20 5-year survival rate of 5% [5].=20 Patients with high tumor microvascular densities have shorter=20 postoperative survival durations than do patients with low = microvascular=20 densities, suggesting that tumor vasculature is important to brain = tumor=20 growth [6,7].=20 Currently, the majority of the available literature data on vessel = formation in brain tumors comes from studies of GBMs. In this = minireview,=20 we describe the characteristics of brain tumor angiogenesis and = the=20 current status and known mechanisms of antiangiogenic therapy for = brain=20 tumors based on published results mainly from studies of primary = brain=20 tumors.

=

=20

Angiogenesis in brain tumors

=20

=20

Tipping the balance between = proangiogenic and=20 antiangiogenic factors

When a solid tumor such as a brain tumor grows = larger than a=20 critical size (1=962 mm in diameter), it must recruit new = blood vessels=20 to have the oxygen and nutrition supply necessary for its survival = and=20 growth. This tumor-induced formation of new blood vessels occurs = primarily=20 via angiogenesis, the process of development and growth of new = blood=20 vessels from pre-existing vasculature [8].

Researchers now widely accept that angiogenesis is = tightly=20 controlled by a balance of pro- and antiangiogenic factors [9=9611].=20 These molecules can be secreted by cancer, endothelial, stromal = and blood=20 cells and the extracellular matrix [12,13].=20 Proangiogenic factors include vascular endothelial growth factor = (VEGF),=20 acidic fibroblast growth factor, basic fibroblast growth factor, = placental=20 growth factor, angiopoietin-2 and interleukins, whereas = antiangiogenic=20 factors include angiostatin, endostatin, thrombospondin 1 and = endothelial monocyte-activating polypeptide 2 [14,15].=20 In addition, the enzymes serine proteinase and metalloproteinase = degrade=20 the extracellular matrix, which has an important role in both the=20 induction and suppression of angiogenesis [16].=20 This biological process, which is essential to not only tumor = development,=20 but also normal development and wound repair, is highly regulated. = When=20 the expression of proangiogenic molecules is balanced with that of = antiangiogenic molecules, the 'angiogenic switch' remains off. = However, in=20 tumor angiogenesis, the tight regulation of the balance of = expression of=20 these molecules is disrupted. Induced expression of proangiogenic=20 molecules leads to uncontrolled and disorganized promotion of = angiogenesis=20 [8,17].

=20

=20

Extracellular signaling promotes = brain tumor=20 angiogenesis

Angiogenesis in solid tumors, including brain = tumors, is=20 believed to be triggered by low oxygen concentrations (hypoxia) = resulting=20 from deficits in the blood supply caused by the tumor's fast = growth.=20 Exposure of brain tumor cells to hypoxia induces expression of=20 hypoxia-inducible factor-1, a transcription factor that regulates = the=20 expression of many angiogenesis- and glucose metabolism-related = genes.=20 Hypoxia-inducible factor-1 activates the transcription of VEGF and = other=20 proangiogenic factors in gliomas, in particular [18=9620].=20 Researchers have found high levels of expression of VEGF mRNA in = the=20 hypoxic regions of high-grade but not low-grade gliomas. In = addition, the=20 VEGF receptors (VEGFR1 and VEGFR2) are highly expressed in gliomas = [21].=20 Expression of VEGF is correlated with microvascular density in = gliomas and=20 meningiomas [22].=20 basic fibroblast growth factor is a potent mitogen of endothelial = cells=20 and is required for glioma angiogenesis in vivo=20 [23].

Signaling by neurotrophins and their receptors = supports=20 neuronal proliferation, differentiation and synapse formation. The = neurotrophin family consists of four structurally related = proteins: nerve=20 growth factor, brain-derived neurotrophin factor, neurotrophin-3 = and=20 neurotrophin-4 [24].=20 Nerve growth factor, brain-derived neurotrophin factor and = neurotrophin-3=20 bind primarily to the receptor kinases TrkA, TrkB and TrkC, = respectively,=20 to mediate their effects across the cell membrane [25].=20 Also, nerve growth factor and brain-derived neurotrophin factor = enhance=20 endothelial cell survival and proliferation [26=9629].=20 In particular, brain-derived neurotrophin factor can enhance the=20 expression of proangiogenic factors (e.g. VEGF) in brain = tumor-derived=20 cells through induction of hypoxia-inducible factor-1 expression = [30].

Interleukin-8 (also known as CXCL8) is a chemokine = with=20 proangiogenic activity. Authors have reported high levels of = expression of=20 hepatocyte growth factor/scatter factor and interleukin-8 in = primary and=20 recurrent glial tumors [31,32].=20 Expression of another chemokine peptide, CXCL12, and its cognate = receptors=20 is induced in brain tumors and promotes angiogenesis [33].=20 In addition, a subset of integrins mediates endothelial-cell = spread and=20 migration in response to growth factor signaling in brain tumor=20 angiogenesis [34].=20 mRNA expression profiles in gliomas from patients have shown = expression of=20 many proangiogenic factors including insulin-like growth factor-1 = in those=20 tumors [35].=20 Stem cell factor and its receptor c-Kit pathway play important = roles in=20 tumor-induced angiogenesis in the brain, as well [36].

=20

=20

γ-Secretase in brain tumor = angiogenesis

Signaling by the transmembrane protein Notch and = its ligand=20 Jagged/Delta is indispensible for neural system development and is = related=20 to the development of many types of tumors [37].=20 Notch signaling is activated by VEGF signaling and suppresses = angiogenesis=20 [38=9640].=20 Accordingly, researchers found that blockade of Delta-like = ligand 4=20 led to increased blood vessel sprouting in a glioma model [41].=20 Interestingly, such increased vessel sprouting does not support, = but=20 rather suppresses, tumor growth, suggesting that Notch signaling = is=20 required for the negative feedback and fine-tuning of the = proangiogenic=20 VEGF signaling to establish functional vessels in brain tumors [41].=20 Notch signaling also downregulates the expression of VEGFR2 and = VEGF in=20 endothelial cells [42].=20 Notch signaling is mediated by cleavage of the Notch molecule by=20 γ-secretase, a presenilin-dependent protease complex [43].=20 VEGF increases γ-secretase activity-mediated Notch 1 = cleavage in=20 endothelial cells. Inhibition of γ-secretase activity blocks = VEGF-induced=20 endothelial cell proliferation, migration and survival, and = eventually=20 leads to decreased angiogenesis [44].=20 In addition, presenilin cleaves the erythroblastic leukemia viral = oncogene=20 homologue 4, ErbB-4 [45],=20 which is widely expressed in gliomas and medulloblastomas and = enhances=20 tumor angiogenesis [46].=20 Moreover, γ-secretase cleaves VEGFR1 [47]=20 and insulin-like growth factor-1 receptor, and both of these = receptors'=20 signaling promotes angiogenesis in astrocytomas and glioblastomas = [35,48].=20 These results suggest that γ-secretase has important complex, = but as yet=20 unidentified, roles in brain tumor angiogenesis.

=20

=20

Intracellular machinery of brain = tumor=20 angiogenic signaling

As described above, researchers have made = considerable=20 progress in understanding the interactions among cell surface = receptors=20 and ligands that regulate angiogenesis. However, the intracellular = machinery that governs signaling from the receptors on the cell = surface to=20 the nucleus to control the induction of angiogenesis remains = poorly=20 understood. Signaling of VEGFR and that of other receptor tyrosine = kinases, such as platelet-derived growth factor receptors and = epidermal=20 growth factor receptors, have regulatory mechanisms that are = similar in=20 many aspects [49].=20 VEGFR signaling may induce activation of Ras/Raf/mitogen-activated = protein=20 kinase [50,51]=20 or phospholipase C-γ/protein kinase C signaling [52],=20 which regulates endothelial cell proliferation, migration and = permeability=20 [53].=20 Also, one of the important signaling pathways activated by VEGFR = is the=20 phosphatidylinositol-3 kinase/phosphatase and tensin=20 homologue/Akt/mammalian target of rapamycin = (PtdIns3K/PTEN/Akt/mTOR)=20 pathway. This PtdIns3K/PTEN/mTOR pathway regulates endothelial = cell=20 survival, translation and permeability [53=9656].=20 This pathway is also activated by other proangiogenic stimuli, = including=20 platelet-derived growth factor, neurotrophins, insulin-like growth = factor,=20 epidermal growth factor and integrins, and plays a critical role = in brain=20 tumor angiogenesis [57].=20 The pivotal role of this signaling pathway in the proliferation = and=20 survival of brain tumor cells strongly suggests the potential use = of=20 inhibitors of it to target both brain tumor cells and blood vessel = endothelial cells [57].

=20

=20

Characteristics of brain tumor = vasculature

=20

=20

The blood=96brain barrier in brain = tumors

The vasculature in a healthy central nervous = system (CNS)=20 tissue is highly specialized and distinguished from the = vasculature in=20 other tissues by a unique structure of blood capillaries, the = blood=96brain=20 barrier (BBB) [58].=20 Unlike other tissues, in which relatively free diffusion of = materials in=20 the blood is allowed through their peripheral capillary walls, the = transportation of materials in the blood circulation to the = peripheral=20 tissues in CNS is tightly regulated by this barrier. The BBB is an = anatomical and physiological barrier that strictly restricts the=20 permeability of blood vessels, suppressing the diffusion of ions,=20 peptides, amino acids and other substances from the bloodstream to = the=20 neural system, while supplying the brain with the required = nutrients for=20 proper CNS function. This barrier is composed of the walls of = vessel=20 endothelia, which are sealed by tight junctions between = endothelial cells.=20 Also, the BBB is wrapped with specialized cells (pericytes) and = the=20 flattened 'endfeet' of astrocytes. Pericytes are relatively=20 undifferentiated mesenchyme-like cells that support capillary = blood=20 vessels. Astrocytes induce the tight junctions of BBB through = decreasing=20 VEGF expression and stimulating angiopoietin release [59].=20 This tight junction prevents the passive diffusion of hydrophilic=20 molecules whose molecular mass is > 500 kDa from the=20 bloodstream to the brain parenchyma. Furthermore, water-soluble = materials=20 are not able to pass through the lipophilic membranes of = endothelial=20 cells. Thus, the BBB is often the primary obstacle to drug = delivery to the=20 CNS [58].

Because the brain is located in a confined space, = leakage of=20 fluid into the brain caused by breakage of the BBB results in = increased=20 interstitial pressure within the skull and, consequently, = vasogenic brain=20 edema. Therefore, keeping the molecules carried in the blood = vessels in=20 the brain from breaching the BBB and entering brain tissue is = essential=20 for maintaining normal brain physiology. Hence, BBB breakdown is=20 associated with many CNS-associated pathologies, including = neurological=20 diseases such as Alzheimer disease.

The BBB in brain tumors is structurally and = functionally=20 abnormal [60=9663].=20 Nevertheless, some features of the normal BBB are retained in = brain tumor=20 vasculature [61,63,64].=20 Researchers found that, in a murine brain tumor metastasis model, = the=20 integrity of the BBB was conserved in small tumors = (< 0.25 mm=20 in diameter) but not in larger tumors [65].=20 In addition to loss of BBB integrity, blood vessels in brain = tumors=20 exhibit abnormal features similar to those in the vessels in other = types=20 of tumors. For example, tumor blood vessels are tortuous, = disorganized and=20 highly permeable because of abnormalities in their endothelial = walls [61=9663,66=9669].=20 Therefore, disruption of the BBB and further increases in the = permeability=20 of tumor blood vessels in brain tumors, because of their loosened=20 endothelial structures, result in increased accumulation of fluid=20 peritumorally and in the surrounding brain, and bring about = vasogenic=20 brain edema. Vasogenic edema is a major cause of morbidity in = patients=20 with brain tumors [70,71].=20 Hence, tumor angiogenesis must be treated properly to not only = prevent=20 brain tumor growth, but also suppress the pathological damage = caused by=20 changes in the permeability of brain tissue. The blood vessels = associated=20 with brain metastases are dilated and contain highly mitotic = endothelial=20 cells [65],=20 which may require high concentrations of VEGF for their growth. = Increased=20 leakage from blood vessels in brain tumors causes suppressed and = irregular=20 blood flow and leads to heterogeneous and inefficient delivery of = oxygen,=20 nutrients and drugs to the brain tumor via the bloodstream [63,71,72].

One of the main causes of increased permeability = and loss of=20 BBB integrity in brain tumor blood vessels is increased expression = of VEGF=20 by the brain tumor cells. Investigators initially purified VEGF = for its=20 ability to induce vascular leakage and permeability, as well as = for its=20 role as a mitogenic factor for endothelial cells. Therefore, it = was=20 originally known as vascular permeability factor as well as VEGF = [73=9675].=20 The effects of VEGF on vascular permeability in the peripheral = circulation=20 appear to occur via modulation of calcium influx, nitric oxide, = activation=20 of guanylyl cyclase, protein kinase G, vesiculo-vacuolar = organelles=20 or increased synthesis of platelet-activating factor in = endothelial cells=20 [73=9678].

Hypoxia and acidosis are hallmarks of solid = tumors, but they=20 are not necessarily correlated with each other [79],=20 and low oxygen and pH levels independently upregulate VEGF = transcription=20 in brain tumors in vivo [60].=20 Also, many oncogenes, such as ras and src, cytokines and growth factor receptor = signaling, and=20 tumor suppressor genes, such as trp53, = regulate=20 transcriptional and translational expression of VEGF in tumors [80,81]=20 and surrounding tissues [12].=20 In principle, tumor blood vessels, especially brain tumor blood = vessels,=20 are abnormal, which means they are highly permeable and even leaky = at many=20 points, resulting in irregular and inefficient blood flow through = them.=20 This irregularity and inefficiency are strongly linked with the = action of=20 VEGF, the major proangiogenic factor.

=20

=20

Targeting brain tumor = vasculature

=20

=20

Therapeutic antiangiogenic agents = for brain=20 tumors

In the early 1970s, Folkman [9]=20 proposed blockage of tumor vascularization as an approach to = treating=20 cancers. Because of the known pivotal role of VEGF in = angiogenesis, many=20 antiangiogenic approaches have targeted VEGF and VEGFR signaling. = More=20 than 30 years after the seminal findings by Folkman's group, = the U.S.=20 Food and Drug Administration approved bevacizumab, a humanized mAb = against=20 VEGF, as the first drug used for antiangiogenic therapy for cancer = (specifically, colon, lung and breast cancer). Also approved for=20 antiangiogenic therapy for renal carcinoma and other tumors by the = U.S.=20 Food and Drug Administration were sorafenib and sunitinib, two = small=20 molecules that target VEGFR kinase activity. Unfortunately, none = of these=20 agents are currently approved as therapy for brain tumors. = However, the=20 number of clinical trials examining the use of antiangiogenic = agents to=20 treat brain tumors is increasing. Table 1=20 summarizes current (as of January 2009) clinical trials targeting=20 angiogenesis in primary and secondary adult brain tumors. More = than 70=20 clinical trials using ∼ 20 anticancer agents that may = inhibit=20 angiogenesis in brain tumors are in progress. The majority of = these trials=20 are targeting VEGF signaling with mAbs against VEGF [20,68],=20 small-molecule tyrosine kinase inhibitors (TKIs) that inhibit = VEGFR2=20 tyrosine kinase activity [82]=20 and soluble decoy receptors developed from VEGFR1 that selectively = inhibit=20 VEGF activity [83].=20 In addition to VEGFR signaling, platelet-derived growth factor = receptor,=20 αvβ3 integrins [84]=20 and intracellular mediators of angiogenic signaling (protein = kinase C and=20 mTOR) are targets of reagents used in these clinical trials. Also, = several=20 clinical trials are studying modulations of BBB integrity designed = to=20 modify the permeability of the blood vessels in brain tumors and=20 eventually improve delivery of cytotoxic agents to the brain = tumors.

=20

Table 1. = Clinical=20 trials with antiangiogenic agents targeting brain=20 tumors^a. bFGF, basic fibroblast growth = factor;=20 GBM, glioblastoma multiform; GS, gliosarcoma; MG, malignant = glioma; OG,=20 oligoglyoma; OA, oligoastrocytoma; MB, medulloblastoma; NB, = neuroblastoma;=20 Meta, metastasized secondary brain tumors; mAb, monoclonal = antibody;=20 PDGFR, platelet-derived growth factor receptor; PKC, protein = kinase C;=20 SKI, serine-threonine kinase inhibitor; PtdIns3K, = phosphatidylinositol-3=20 kinase; FKBP-12, FK506 binding protein 1A- 12 kDa.=20

Drug	Target	Phase	Tumors

XL184 (TKI)	VEGFR, MET	II	GBM
ZK222584 (TKI)	VEGFR, PDGFR	I,II	MG
Pazopanib (TKI)	VEGFR/PDGFR/c-Kit	II	Glioma, Meta
Sorafenib (TKI)	VEGFR/PDGFR/Raf	I,II	GBM, Meta,GS
Sunitinib (TKI)	VEGFR/PDGFR	I,II	GBM,Meta
Thalidomide/ = Lenalidomide	VEGF/bFGF/TNF	I,II	GBM,Meta
VEGF Trap (Peptide)	VEGF	II	MG
Ceradinib (TKI)	VEGFR	I,II,III	GBM
ZD6474 (TKI)	VEGFR	I,II	GBM
CT-322 (Peptide)	VEGFR	I,II	GBM
Bevacizumab (mAb)	VEGF	I,II	GBM
Valproic acid	Potential anti-angiogenic	II	Meningioma, GBM
Cilengitide (Peptide)	Integrin	II,III	GBM, MG
Everolimus/ = Sirolimus	FKBP-12/mTOR	I, II	MG

II	GBM
RAD001 (SKI)	mTOR	III	Astrocytoma
Temsirolimus (SKI)	mTOR	I	GBM
Tandutinib (TKI)	PDGFR	I,II	GB,Astrocytoma,Meta
XL765 (SKI)	PtdIns3K/mTOR	I	Oligodedroglioma
Enzastaurin (SKI)	PKC	I,II	GBM
MPC-6827	Microtubule (microvessel) =	I,II	GBM
CY997	tubulin (microvessel)	I,II	GBM, Gliosarc, = Meta
Sodiumthiosulfate/Mannitol	BBB	I,II	OG,OA,MB,NB,GBM

^a The = information in=20 this table was extracted from the National Cancer = Institute=20 Web site (http://www.cancer.gov/) in a search = for=20 currently (January 2009) open clinical trials. Most of = the=20 drugs have been used in multiple concurrent clinical=20 trials.

Because Notch signaling suppresses and = orchestrates the=20 formation of brain tumor vessel sprouts, treatment with = γ-secretase=20 inhibitors may inhibit normal Notch signaling and thus tumor = angiogenesis.=20 Indeed, the authors reported that treatment with a = γ-secretase inhibitor=20 suppressed the growth and vascularization of human GBM xenografted = into=20 nude mice [85].

The intracellular machinery of angiogenic = signaling=20 described above can be an effective target in antiangiogenic = therapy for=20 brain tumors. As expected from the observation that the = PtdIns3K/Akt/mTOR=20 signaling pathway is essential for endothelial cell survival and = blood=20 vessel permeability, inhibition of mTOR signaling by = small-molecule=20 inhibitors or RNA interference have proven to be efficacious at=20 antiangiogenic therapy in preclinical models of malignant glioma = [86,87].=20 The signalling in brain tumor angiogenesis and its targeting are=20 summarized in Fig.=20 1.

=20

= Fig. 1.=20 Angiogenic interaction between brain tumor and=20 endothelial cells and antiangiogenic therapy. VEGF = expression is=20 induced in tumor cells by hypoxia [19],=20 radiation, receptor tyrosine kinase signaling and Akt = pathways [114].=20 VEGF and other proangiogenic factors are secreted by brain = cancer=20 cells in a paracrine and endocrine manner [83].=20 These factors support endothelial cell survival. Also, VEGF = induces=20 permeability of the BBB [78].=20 Notch signaling mediated by γ-secretase suppresses = angiogenesis but=20 is required for proper vascular development [38=9640,44].=20 The signaling required for angiogenesis in brain tumors can = be=20 targeted using an antibody against VEGF [20,68],=20 a decoy receptor to VEGF [115],=20 small-molecule TKIs of VEGFR[82],=20 inhibitors of other kinases in the signaling cascade and = inhibitors=20 of γ-secretase [85].=20 Ab, antibody; HGF/SF, hepatocyte growth factor/scatter = factor; IR,=20 ionizing radiation; RTK, receptor tyrosine kinase; PTEN, = phosphatase=20 and tensin homologue; PI3K, phosphatidylinositol-3 kinase; = HRE,=20 hypoxia-responsive element; ROS, reactive oxygen species; = γ-Sec,=20 γ-secretase.

[Normal View=20 ]

=20

=20

Efficacy of antiangiogenic therapy = for brain=20 tumors

The results of the first series of clinical trials = (phase I=20 and II) on treatment of GBM with vatalanib, a small-molecule TKI = against=20 of VEGFR2, were disappointing [88,89].=20 However, recent clinical trials showed that bevacizumab and a TKI = of VEGFR=20 were effective in the treatment of brain tumors when combined with = standard chemotherapeutic agents [20,82].=20 Researchers in one of these studies observed a strong antiedema = effect of=20 bevacizumab, suggesting that this antiangiogenic therapy decreased = the=20 permeability of brain tumor blood vessels [20].=20 These results suggested that the antiangiogenic agents used = augmented=20 cytotoxic drug efficacy by improving blood flow in these tumors = [20,82].=20 The novel hypothesis that 'antiangiogenic therapy normalizes the = abnormal=20 tumor blood vessels' may explain these results [90].=20 As described above, intratumoral vessels are leaky because of high = local=20 concentrations of VEGF, a potent vascular permeability factor. = Blocking=20 VEGF or VEGFRs in tumors suppresses the function of VEGF = signaling; the=20 tumor vasculature permeability then returns to normal, leading to = the=20 restoration of effective blood flow [91].=20 Accordingly, antiangiogenic therapy enhances the delivery and = efficacy of=20 concurrently administered cytotoxic agents [90,92=9694].=20 This is supported by the results of studies using a number of = preclinical=20 models [63,95=9697],=20 including primary and secondary brain tumor models [68,98].=20 Authors also reported that VEGFR2 blockade by the mAb DC101 = temporarily=20 normalized tumor vessel walls, improving vascular function and = tumor=20 oxygenation and thus enhancing responses to radiotherapy in a GBM = model in=20 nude mice [99].=20 In human clinical trials, treatment with bevacizumab indeed = changed=20 several abnormal characteristics of tumor vessels to resemble the = normal=20 characteristics of non-tumor vessels [100].=20 In patients with recurrent GBM treated with a novel TKI, the VEGFR = inhibitor AZD2171, the treatment normalized tumor blood vessels, = leading=20 to the significant clinical benefit of alleviation of edema [82].=20 However, such normalization seems to be only temporal and = reversible,=20 suggesting that the optimal window of time during which combined = cytotoxic=20 and antiangiogenic therapy for brain cancer provides the best = outcome is=20 limited. It has been reported that during this window, after = treatment=20 with VEGFR2 antibody, tumor oxygenation improved and pericyte = coverage and=20 angiopoietin expression increased and thus decreased interstitial = pressure=20 and vessel permeability in orthotopic GBM [68].

Researchers have suggested that antiangiogenic = therapeutic=20 agents have several mechanisms of action in combined treatment = with=20 radiotherapy [101]=20 For example, a study found that blocking VEGF's action enhanced = the=20 cytotoxic effect of radiotherapy for tumor cells in vitro [102].=20 Also, authors reported that the response of brain tumors to = radiotherapy=20 depended on endothelial cell apoptosis in the tumors in a murine = model [103].=20 This suggested that blockage of VEGF signaling leading to = apoptosis of=20 endothelial cells and thus enhanced the efficacy of radiotherapy.=20 Radiotherapy also induced expression of hypoxia-inducible factor-1 = and=20 genes downstream of it, including VEGF, in an in vivo murine tumor model [104].=20 In brain tumor models, radiotherapy has induced VEGF expression; = this=20 induced expression is essential for endothelial cell survival [105=96107].=20 Therefore, radiotherapy-induced endothelial cell death would be = enhanced=20 by blockage of VEGF action by inhibition of newly expressed VEGF = in=20 irradiated cancer cells and endothelial cells.

Compelling evidence suggests that cancer cells are = generated=20 by small fractions of self-renewing, multipotent, tumor-initiating = cells=20 termed cancer stem cells (CSCs) in brain tumors as they are in = other=20 tumors [108].=20 Calabrese et al. [109]=20 observed that brain tumor CSCs are found in vascular niches in the = tumors=20 as normal brain stem cells are. These authors reported that = antiangiogenic=20 therapy disrupted the tumor vasculature and that the CSC niche=20 microenvironment associated with the tumor blood vessels reduced = the CSC=20 population in the brain, suggesting that extirpation of brain CSCs = may=20 contribute to the efficacy of antiangiogenic cancer = therapy.

=20

=20

Current limits of and perspectives = on=20 antiangiogenic therapy for brain tumors

The use of antiangiogenic therapy for brain tumors = carries=20 several concerns [110].=20 Despite angiogenesis being essential for the growth of all tumors, = antiangiogenic therapy usually results in only transitory delays = in tumor=20 growth and progression, after which tumors begin to regrow and the = cancer=20 progresses. This may result from the adaptation of tumors to=20 antiangiogenic therapy; evasive resistance [111]=20 and intrinsic resistance of tumors to antiangiogenic therapy. = Tumor cells=20 self-adjust to antiangiogenic therapy by developing alternative = pathways=20 to recruit new blood vessels. Also, tumors may activate and = enhance=20 invasion and metastasis of their own cells after antiangiogenic = therapy=20 [111].=20 Therefore, use of antiangiogenic therapy combined with other = therapies=20 that may override this adaptive 'self-adjusting' pathway would=20 significantly improve the efficacy of the anticancer = treatment.

Hemorrhage and thrombosis are potential = complications of=20 antiangiogenic treatment of GBM [20].=20 Also, authors have reported intracerebral hemorrhages in patients = with=20 gliomas treated with bevacizumab in combination with irinotecan, a = topoisomerase inhibitor [112].=20 Rebound cerebral edema caused by the reversibility of = normalization of=20 blood vessels in brain tumors after stopping antiangiogenic = treatment is=20 another potential problem. In addition, blockage of angiogenesis = is=20 potentially cytotoxic to neural stem cells, which reside in = perivascular=20 niches in the subventricular zones of the brain and are supported = by=20 growth factors generated by the endothelial cells. Therefore,=20 antiangiogenic therapy resulting in VEGF-signaling suppression or=20 endothelial cell death may cause neural stem cell death [113].

When more clinical data on the efficacy and = mechanism of=20 action of antiangiogenic therapy for brain and other tumors from = many of=20 the ongoing clinical trials of this therapy become available, the=20 resulting improvement in understanding of this mechanism will = allow us to=20 optimize combinations of antiangiogenic therapy and other = cytotoxic=20 treatments and improve the benefits of this therapy in patients = with brain=20 cancer.

=20

=20

Addendum

After this article was submitted, U.S. Food and = Drug=20 Administration (FDA) granted accelerated approval of bevacizumab = (antibody=20 to VEGF) for glioblastoma patients with progressive disease = following=20 prior therapy, in May 2009. This is the first FDA approved = anti-angiogenic=20 agent for brain tumors.

=20

=20

=20

Acknowledgements

This work was supported by National Institutes of = Health=20 grants R01 CA109520 and CA100816-01A1 (to H.-Y. Lee).

=20

=20

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