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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_01C9F71F.4E97FC40 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_0000_01C9F71F.4E97FC40 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://download.interscience.wiley.com/freeflow/js/site.js /************************************************************************= ***************************=0A= FILE: site.js=0A= AUTHOR: Jonathan Sugar, Chris Neatby-Smith and Digital Pulp. July = 2003.=0A= =0A= DESCRIPTION: Javascript relating to site wide functionality across =0A= InterScience.=0A= =0A= INCLUDES FUNCTIONS=0A= =3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=0A= =0A= Search Functions=0A= ----------------=0A= =0A= selectAndSubmit() - for deciding where to submit a query based on the = state of the form.=0A= processallContentSearch() - for validating form input for an all = content search.=0A= processTitleSearch() - for validating form input for a Title search.=0A= checkCitationSearch() - for validating the citation search form.=0A= selectAll() - check all the checkboxes in a form.=0A= do_popup() - stasa Acronym Finder window popup function.=0A= =0A= =0A= Other Functions=0A= ---------------=0A= =0A= fixInlineLists() - fix netscape 4.x bug where

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

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

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

You do not have = anything saved in this menu.

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

" = + liels[j].innerHTML + "

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

" + liels[j].innerHTML + "

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

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Cancer Science

ISSN: 1349-7006 (Online)

ISSN: 1347-9032 (Print)

Volume 9999, Issue 9999, 2009.

Pages: ???=96???

Top of Page
Abstract
Materials and Methods
Results
Discussion
Acknowledgments
Disclosure Statement
References

List of=20 Images

Abstract=20 Only

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3D"About

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

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 © 2009 Japanese = Cancer Association

'); } =0A= function imagelink(hwnd, iname) {if (iname.indexOf(' ') > -1) {var = images =3D iname.split(' ');for (var i =3D 0; i < images.length; ++i) = {hwnd.document.write('

');}} = else {hwnd.document.write('

');}}=0A= function otherlinks(hwnd, iname, size) { hwnd.document.write('View List of Images = '); hwnd.document.write('

'); } =0A= =0A= =0A= function caption_f1(hwnd) = {hwnd.document.write('Fig. 1. shRNA-mediated = knockdown of tenascin-C does not affect cell proliferation in LN229 = glioblastoma cells. (a) Lentiviral shRNA against control and tenascin-C = was introduced into LN229 cells and IRES-mediated EGFP expressions were = seen as an infection marker. Fluorescence (EGFP) and phase-contrast (PH) = images are shown. (b) Representative immunoblot of tenascin-C and = α-tubulin in shRNA-mediated protein knockdown against control and = tenascin-C in LN229 cells. (c) Quantification of cell growth in control = and tenascin-C knockdown cells 2 and 4 days after cell plating. = Data are shown as mean ± SD (n =3D 3). = Scale bar, 200 µm; TNC-KD, tenascin-C-knockdown cells.'); }=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= function caption_f2(hwnd) = {hwnd.document.write('Fig. 2. Endogenous = tenascin-C regulates cell motility in LN229 glioblastoma cells. (a) The = confluent monolayers of the control and tenascin-C-knockdown LN229 cells = were photographed 0 and 14 h after scraping. (b) The distance of = cell-edge movements 14 h after scraping is shown as = mean ± SD (n =3D 3). (c) F-actin = (phalloidin) and tyrosine-phosphorylated proteins were stained in the = control and tenascin-C-knockdown cells on the non-coated and fibronectin = coated coverslips. (d,e) The average area and intensity of = tyrosine-phosphorylation signals in the control and tenascin-C knockdown = cells stained by phosphor-tyrosine antibody on the non-coated = glass-bottom dishes are measured and shown as mean ± SE = (control, n =3D 30; TNC-KD, n =3D 36). = (f) Representative immunoblot of phosphorylated p130Cas and FAK in = shRNA-mediated protein knockdown against control and tenascin-C in LN229 = cells. TNC-KD, tenascin-C-knockdown cells; **, = P < 0.001; scale bar, 200 µm.'); }=0A= function display_f2(size) { var i =3D new Array(); i[0] =3D 'nf2'; i[1] = =3D 'nf2'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f2', size); imagelink(hwnd, i[size]); = caption_f2(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f3(hwnd) = {hwnd.document.write('Fig. 3. Endogenous = tenascin-C activates glioblastoma cell locomotion. Cell tracking of (a) = control cells, (b) tenascin-C-knockdown cells and (c) = tenascin-C-knockdown with addition of exogenous tenascin-C. Cells were = imaged every 5 min for 12 h under microscopy. (d) Track length = (µm). (e) Mean velocity (µm/min). (f) Distance from the = origin. Tracks and velocities were calculated with MetMorph software = (Molecular Devices, Sunnyvale, CA). TNC-KD, tenascin-C-knockdown cells; = TNC-KD + TNC, tenascin-C-knockdown cells with exogenous = tenascin-C; *, P < 0.01; **, = P < 0.001; scale bars, 50 µm.'); }=0A= function display_f3(size) { var i =3D new Array(); i[0] =3D 'nf3'; i[1] = =3D 'nf3'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f3', size); imagelink(hwnd, i[size]); = caption_f3(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f4(hwnd) = {hwnd.document.write('Fig. 4. Endogenous = tenascin-C enhances glioblastoma invasion with reactive change of the = brain tissue in a xenograft glioblastoma model. Intracranial tumors = derived from (a,b,c,d) control cells and (e,f,g,h) tenascin-C-knock down = cells are shown by hematoxylin–eosin staining (HE), IRES-mediated = EGFP expression of implanted cells (EGFP) and tenascin-C staining (TNC). = Small and large rectangles in (a) and (e) are enlarged in (b,f) and = (c,d,g,h), respectively. Small white rectangles are enlarged to large = white rectangles in (c,d). (i) Tumor size is shown as = mean ± SD (n =3D 6). (j) Number of = cell clusters per circumference of the tumor core is shown as = mean ± SD (n =3D 6). Scale bars, = 1 mm (a,e), 100 µm (b,f), 500 µm (c,d,g,h); = *, P < 0.01.'); }=0A= function display_f4(size) { var i =3D new Array(); i[0] =3D 'nf4'; i[1] = =3D 'nf4'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f4', size); imagelink(hwnd, i[size]); = caption_f4(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f5(hwnd) = {hwnd.document.write('Fig. 5. Correlation between = tenascin-C expression and reactive change of peritumoral brain = parenchyma in resected glioblastomas. (a) Representative MR images of = glioblastoma show the tumor core enhanced by gadolinium (Gd enhanced) = and the peritumoral change of brain parenchyma as hyperintense area of = FLAIR sequence. (b) Correlation between FLAIR/Enhanced volume ratio and = tenascin-C mRNA expression in 16 patients with glioblastoma. (c) = Correlation between proliferation activity (Ki-67 index) and tenascin-C = expression in 16 resected glioblastomas. a. u., arbitrary unit; = r, Spearman correlation coefficient.'); }=0A= function display_f5(size) { var i =3D new Array(); i[0] =3D 'nf5'; i[1] = =3D 'nf5'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f5', size); imagelink(hwnd, i[size]); = caption_f5(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f6(hwnd) = {hwnd.document.write('Fig. 6. Infiltrating = glioblastoma cells show cytoplasmic deposition of tenascin-C in reactive = changes of peritumoral brain parenchyma detected by FLAIR image. = Representative images of the tumor core and peritumoral tissue in = patients with (a) low FLAIR/Enhanced volume ratio (F/E ratio) and (b) = high F/E ratio. Each square in MR images (left panel) indicates the = localization of the microscopic images (three right panels; HE, Ki-67, = TNC). Hematoxylin–eosin staining (HE), mitotic activity staining = (Ki-67), tenascin-C staining (TNC). Scale bars, 200 µm.'); }=0A= function display_f6(size) { var i =3D new Array(); i[0] =3D 'nf6'; i[1] = =3D 'nf6'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f6', size); imagelink(hwnd, i[size]); = caption_f6(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('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');hwnd.document.write('

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Published Online: 20 Apr 2009

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Abstract | References | Full Text: = HTML , PDF (5084k)=20

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3D""=20

3D""=20

Endogenous tenascin-C enhances = glioblastoma=20 invasion with reactive change of surrounding brain = tissue

Eishu=20 Hirata ^1,2 = ,=20 Yoshiki Arakawa ^1,5 = , Mitsuaki Shirahata ¹ = , Makoto Yamaguchi ¹ = , Yo Kishi ¹ , = Takashi Okada ¹ , = Jun A. Takahashi ³ = , Michiyuki Matsuda ² = and Nobuo Hashimoto ⁴

¹ Department of=20 Neurosurgery , ² = Department of=20 Pathology and Biology of Diseases, Kyoto University Graduate = School of=20 Medicine, Kyoto 606-8507 ; ³ Department of = Neurosurgery,=20 Kitano Hospital, Osaka 530-8480 ; ⁴ National = Cardiovascular=20 Center, Osaka 565-8565, Japan

⁵ To whom = correspondence=20 should be addressed. E-mail: yarakawa@sb3.so-net.ne.jp=20

Copyright =A9 2009 Japanese Cancer=20 Association

ABSTRACT

3DAbstract=20

Tenascin-C is an extracellular matrix glycoprotein = implicated in embryogenesis, wound healing and tumor progression. = We=20 previously revealed that tenascin-C expression is correlated with = the=20 prognosis of patients with glioblastoma. However, the exact role = of=20 endogenous tenascin-C in regulation of glioblastoma proliferation = and=20 invasion remains to be established. We show here that endogenous=20 tenascin-C facilitates glioblastoma invasion, followed by reactive = change=20 of the surrounding brain tissue. Although shRNA-mediated knockdown = of=20 endogenous tenascin-C does not affect proliferation of = glioblastoma cells,=20 it abolishes cell migration on a two-dimensional substrate and = tumor=20 invasion with brain tissue changes in a xenograft model. The = tyrosine=20 phosphorylation of focal adhesion kinase, a cytoplasmic tyrosine = kinase=20 that associates with integrins, was decreased in = tenascin-C-knockdown=20 cells. In the analysis of clinical samples, tenascin-C expression=20 correlates with the volume of peritumoral reactive change detected = by=20 magnetic resonance imaging. Interestingly, glioblastoma cells with = high=20 tenascin-C expression infiltrate brain tissue in an autocrine = manner. Our=20 results suggest that endogenous tenascin-C contributes the = invasive nature=20 of glioblastoma and the compositional change of brain tissue, = which=20 renders tenascin-C as a prime candidate for anti-invasion therapy = for=20 glioblastoma. (Cancer Sci = 2009)

(Received November 25, 2008/Revised March 27, = 2009/Accepted=20 April 6, 2009)

DIGITAL OBJECT IDENTIFIER (DOI)

10.1111/j.1349-7006.2009.01189.x About=20 DOI

=20

=20

Article Text

Glioblastoma is the most common and most agressive = primary=20 brain tumor in adults. As it is recalcitrant to any current = treatment=20 including surgery, radiotherapy and chemotherapy, the median = survival time=20 is less than 1 year.⁽¹⁾= =20 Glioblastoma is characterized by extensive invasiveness into the=20 surrounding brain tissue, which makes it impossible to provide a = cure by=20 surgical resection.⁽²⁾= =20 For these reasons, recurrent tumors develop adjacent to the = resection=20 cavity and in the contralateral hemisphere. Magnetic resonance = (MR)=20 imaging is the standard clinical examination for the diagnosis of=20 glioblastoma. Glioblastoma is a radiologically heterogeneous tumor = with=20 severe edema of the brain parenchyma. Gadolinium enhancement often = demonstrates 'ring enhancement', which represents central necrosis = and=20 enhancing rim of glioblastoma cells, with accompanying = microvascular=20 hyperplasia and increased permeability.⁽³⁾= =20 The fluid-attenuated inversion recovery (FLAIR) sequence yields = heavily=20 T2-weighted MR images of the brain with suppression of the = cerebrospinal=20 fluid signal. Glioblastoma induces vasogenic edema and changes the = composition of brain parenchyma. As a possible mechanism,=20 glioblastoma-derived factors downregulate endothelial tight = junction=20 proteins, resulting in impairment of the blood=96brain=20 barrier.⁽⁴⁾= =20 Various extracellular matrices, such as matrix metalloproteinase = (MMP),=20 collagen type IV and laminin are=20 increased.⁽⁵⁾= =20 For these reasons, FLAIR images can detect the peritumoral change = in brain=20 tissue as hyperintense.⁽⁶⁾=

This invasive capacity of glioma cells is driven = by complex=20 molecular processes involving the remodeling of extracellular = matrix=20 (ECM), intracellular signaling and cytoskeletal=20 reorganization.⁽⁷⁾= =20 Several genes involved in glioma invasiveness have been = identified,=20 including those coding for chemoattractants, signaling molecules, = adhesion=20 molecules and MMP.⁽⁸⁼⁹⁶¹⁰^{)<=
/SUP>=20
Migration of glioma cells involves active intracellular signaling =
with=20
molecules such as PI3 K, focal adhesion kinase (FAK) and =
small GTPase=20
proteins.⁽¹¹^{)=20
For example, the Rho GTPase RhoA and its effector, mDia1, regulate =
cell=20
polarity and focal adhesion turnover in migrating glioma cells by=20
localizing adenomatous polyposis coli (Apc) and =
proto-oncogene-encoded=20
c-Src tyrosine kinase (c-Src).⁽¹²^{)=20
Src binding to FAK contributes to src activation, promoting =
further=20
phosphorylation of FAK at additional tyrosines, which is =
responsible for=20
efficient disassembly of focal =
adhesion.⁽¹³^{)=20
Meanwhile, glioma invasion requires the detachment of the cell =
from the=20
ECMs and/or the adjacent cells as well as proteolytic degradation =
of ECMs=20
around the tumor.⁽²⁾=}}}}

Tenascin-C, a component of the ECMs, is highly = expressed in=20 many malignancies, including = glioblastoma.⁽^14=9617⁾=20 Many previous reports have provided important insights into the = biological=20 function of tenascin-C. However, the mechanisms by which it = signals are=20 not yet clear, and these appear to be variable across cell types = and=20 experimental models.(¹⁴^{)=20
In addition, most studies have relied upon exogenous tenascin-C =
and in vitro models,⁽^18=9622⁾=20
while few have focused upon this protein in =
vivo.=20
Curiously, melanoma invasion in vivo was =
only=20
slightly promoted in knockout mice lacking=20
tenascin-C.(}²³⁾

In this study, therefore, we investigated the role = of=20 endogenous tenascin-C in glioblastoma invasion and proliferation = by using=20 shRNA-mediated knockdown of its expression. To characterize the = function=20 of intrinsic tenascin-C of glioblastoma cells in=20 vivo, we used a glioblastoma xenograft model. In the = analysis of=20 resected tumor in patients with glioblastoma, we examined the = correlation=20 between tenascin-C expression, glioblastoma invasion and secondary = tissue=20 change of brain parenchyma. Our results provide a new insight into = the=20 role of endogenous tenascin-C in glioblastoma = progression.

=20

=20

Materials and Methods

=20

Antibodies and=20 reagents. The following primary and secondary = antibodies were=20 used for Western blot analysis, immunocytochemistry or=20 immunohistochemistry: anti-human tenascin-C mouse monoclonal = antibody=20 (Abcam, Cambridge, MA), anti-human/mouse tenascin-C rat monoclonal = antibody (R&D Systems, Inc., Minneapolis, MN); anti-human = tenascin-C=20 mouse monoclonal antibody (clone 49; Novocastra Laboratories, = Newcastle, UK); anti-human Ki-67 mouse monoclonal antibody=20 (clone MM1, Novocastra Laboratories); anti-human = α-tubulin mouse=20 monoclonal antibody (TU-02; Santa Cruz Biotechnology, Santa Cruz, = CA);=20 anti-phosphotyrosine mouse monoclonal antibody (4G10) (Millipore = Corporate=20 Headquarters, Billerica, MA); anti-phosphotyrosine mouse = monoclonal=20 (P-Tyr-100) anti-phospho-p130 CAS (Tyr165) rabbit polyclonal = antibodies=20 (Cell Signaling Technology, Beverly, MA); anti-p130 CAS, anti-FAK = (clone=20 77) and anti-phospho-FAK (pY397) mouse monoclonal antibodies (BD=20 Bioscience, San Jose, CA); and AlexaFluora 405-goat anti-mouse and = 546-goat anti-rat IgG antibodies (Invitrogen, Carlsbad, CA).=20 Rhodamine-conjugated phalloidin (Invitrogen) was used to stain=20 F-actin.

=20

Cell = culture. LN229 human=20 glioblastoma cells were obtained from the American Type Culture = Collection=20 (Manassas, VA) and maintained in Dulbecco's modified Eagle's = medium (DMEM)=20 containing 10% fetal bovine serum (FBS) at 37=B0C in a humidified = atmosphere=20 containing 5% CO₂. For the evaluation of = cell=20 growth, 5 =D7 10⁴ glioma cells = were seeded=20 onto 24-well plates, and the number of viable cells was counted on = days 2 and 4.

=20

Lentiviral shRNA-mediated = RNA=20 interference. The knockdown vector was constructed by = using a=20 shRNA expression lentiviral vector. Lentiviral production and = infection=20 were described previously.⁽²⁴^{)=20
The target sequence for RNA interference was selected from human=20
tenascin-C candidate sequences that are available from GenomeWide =
siRNA=20
(Qiagen, Hilden, Germany). The selected target sequence for =
tenascin-C=20
corresponded to the mRNA of the coding region, =
nt 3853=963871,=20
5'-GCTGGGAT GCCCTCAAACT-3'. The sequence 5'-GATTCTCCGAACG =
TGTCAC-3' was=20
used as a negative control.}

=20

Fluorescence microscopy = and image=20 analysis. Our method of fluorescence microscopy is = described=20 previously.⁽²⁵^{)=20
Glioma cells were cultured on non-coated and fibronectin-coated=20
glass-bottom dishes in DMEM with 10% FBS at 37=B0C in a humidified =

atmosphere containing 5% CO₂. After =
24 h,=20
cells were fixed with 4% paraformaldehyde, permeabilized with 0.2% =
Triton=20
X-100 in PBS, blocked with 5% BSA and 1% FBS in PBS, and incubated =
with=20
the primary antibody overnight, followed by incubation with=20
fluorophore-conjugated secondary antibodies. After rinsing in PBS, =
the=20
cells were imaged with a confocal laser scanning microscope =
(FV1000;=20
Olympus, Tokyo, Japan). To compare the level of tyrosine =
phosphorylation=20
at the focal adhesion, the area and intensity of the =
phospho-tyrosine=20
signal at focal adhesions were measured and corrected by MetMorph =
software=20
(Molecular Devices, Sunnyvale, CA). The analyses were in the =
control (3980=20
focal adhesions in 30 cells) and the tenascin-C knockdown (5072 =
focal=20
adhesions in 36 cells).}

=20

Western blot=20 analysis. Cultured cells at 80% confluency were washed = with=20 PBS and the cellular proteins were extracted in cell lysis buffer=20 (Invitrogen) supplemented with 2.5% 2-mercaptoethanol and boiled = at 95=B0C=20 for 5 min. Equal amounts of proteins were electrophoresed in = 4=9612%=20 precasted Tris-Bis gels (Invitrogen) and transferred onto = nitrocellulose=20 membranes with the use of the Iblot system (Invitrogen). After = blocking=20 with 5% bovine serum albumin (BSA) in Tris-buffered saline (TBS) = with=20 0.05% Tween 20 (TBS=96T), the membrane was incubated with = primary=20 antibody overnight at 4=B0C, followed by washing in TBS=96T and = incubation=20 with a horseradish peroxidase-conjugated goat anti-mouse IgG = diluted in=20 TBS=96T with 5% skim milk for 1 h at room temperature. The = labeled=20 proteins were visualized with ECL plus Western blot detection = system (GE=20 Healthcare, Piscataway, NJ).

=20

Monolayer wound healing = assay and cell=20 locomotion assay. A wound was made by scratching the = monolayer=20 cells with a pipette tip.⁽¹²^{)=20
After the removal of scratched cells, the cells were cultured for=20
14 h. Photographs of cells were taken before and after =
wounding to=20
measure the width of the wound. For the analysis of single cell=20
locomotion, cells were seeded onto either a non-coated or =
tenascin-C=20
coated (10 =B5g/mL) glass-bottom dish 24 h before =
imaging. Cells=20
were imaged every 5 min for 12 h with an Olympus IX81 =
inverted=20
microscope (Olympus) equipped with the ZDC autofocusing system. =
Cell track=20
and velocity were calculated with MetMorph software.}

=20

Mouse xenograft glioma=20 model. Animal care and experiments complied with = Japanese=20 community standards on the care and use of laboratory animals. For = intracranial transplantation, glioma cells=20 = (5 =D7 10⁵ cells/5 =B5L of = PBS)=20 were stereotactically transplanted into the right striatum of=20 9-week-old-male nude mice (BALB/cAJc1-nu) as described=20 elsewhere.⁽²⁶^{)=20
Neurological defects and emaciation of the mice were carefully =
observed=20
every day. Three weeks after transplantation, the brain specimens =
were=20
prepared after euthanasia. The brain tissues were fixed in 4%=20
paraformaldehyde in PBS. Sectioned coronally at the point of =
cellular=20
implantation, the brain tissue was embedded into Optimal Cutting=20
Temprature (O.C.T.) compound (Sakura, Tokyo, Japan) and frozen in =
liquid=20
nitrogen. Twenty-micrometer sections were cut and mounted on =
silane-coated=20
slides, and observed through a fluorescence microscope system=20
(BX51N-34-FL-1; Olympus).}

=20

Resected glioblastoma = samples,=20 quantitative reverse transcription PCR (RT-PCR) and=20 immunohistochemistry. Histological diagnosis was = conducted=20 according to the WHO Classification of Brain=20 Tumors.⁽¹⁾= =20 Tumor specimens for molecular research were snap-frozen = immediately after=20 surgical resection and kept at =9680=B0C. For the sampling of = tumor tissues=20 corresponding to the gadolinium-enhanced lesion or hyperintense = area of=20 FLAIR sequences, we used either StealthStation TRIA plus with = Cranial v4.0=20 software (Medtronic Sofamor-Danek), or Vector Vision Compact = Navigation=20 System with VV Cranial v7.5 software (BrainLab AG, Heim Stetten, = Germany).=20 The study protocol was approved by the institutional review board = of Kyoto=20 University, and written informed consent was obtained from each = patient.=20 For the immunohistochemical analysis in resected glioblastoma = samples, the=20 conventional avidin=96biotin=96peroxidase complex method was used = as described=20 elsewhere.⁽²⁷^{)=20
For quantitative reverse RT-PCR, we used methods described=20
previously.⁽²⁸^{)=20
The second-derivative maximum method was used for crossing-point=20
determination using LightCycler Software 3.3 (Roche, Basel, =
Switzerland).=20
GAPDH was used as an internal control. The following primer sets =
were=20
used: forward, 5'-GTCACCGTGTCAACCTGATG-3', and reverse,=20
5'-GCCTGCCTTCAAGATTTCTG-3', for tenascin-C; forward,=20
5'-TGACAACAGCCTCAAGATCA-3', and reverse, =
5'-CTGTGGTCATGAGTCCTTCC-3', for=20
GAPDH. The conditions for PCR were 45 cycles of 95=B0C for =
10 s for=20
denaturation, 56=B0C for 5 s for annealing and 72=B0C for =
10 s for=20
extension.}}

=20

Measurement of tumor = volume and=20 definition of 'FLAIR/Enhanced volume ratio'. To = calculate=20 'FLAIR/Enhanced volume ratio' in each patient, we defined two = tumor=20 volumes on the multislice MR images. For measurement of 'Enhanced = tumor=20 volume', the border of the tumor area was determined at the = enhancing rim=20 on the T1-weighted images with gadolinium enhancement. For = measurement of=20 'FLAIR hyperintensity volume', the border of the tumor area was = determined=20 at the outer rim of the hyperintensity area in FLAIR sequences. = The planar=20 dimension was analyzed using NIH Image software. In either case, = tumor=20 volumes were calculated by the formula: Σ([planar = dimension]=D7[slice=20 distance]). 'FLAIR/Enhanced volume ratio' was defined by the = formula:=20 ([FLAIR tumor volume] =96 [gadolinium-enhanced tumor=20 volume])/[gadolinium-enhanced tumor volume]) (%).

=20

Statistical = analysis. The=20 Spearman rank correlation test was used to analyze correlations = between=20 tenascin-C expression and Invasion Index. When two groups were = compared,=20 an unpaired Student's t-test was=20 applied.

=20

=20

Results

=20

shRNA-mediated knockdown = of endogenous=20 tenascin-C does not affect glioblastoma cell=20 proliferation. The biological functions of tenascin-C = have=20 been implicated in tumor progression.⁽²⁹^{)=20
However, relevant studies to date have been performed using =
exogenous=20
tenascin-C. To examine the role of endogenous tenascin-C in =
glioblastoma=20
cells, LN229 cells were transduced with a lentiviral shRNA =
sequence=20
against tenascin-C to knock down this protein's expression. The =
lentiviral=20
transduction efficiencies of tenascin-C shRNA and control shRNA =
were=20
consistently greater than 90% as measured by IRES-mediated =
enchanced-green=20
fluorescent protein (EGFP) expression (Fig. 1a).=20
Tenascin-C is highly expressed in LN229 cells, and the shRNA for=20
tenascin-C significantly reduced its protein expression (Fig. 1b).=20
Although tenascin-C has been shown to mediate its effects on cell=20
proliferation via mitogen-activated protein kinase signaling in =
cells=20
plated on fibronectin,⁽²⁰^{)=20
there was no significant difference in the growth rates between =
the=20
control shRNA and the tenascin-C-knockdown glioblastoma cells on =
the=20
substrata without fibronectin (Fig. 1c).=20
This observation indicates that endogenous tenascin-C does not =
accelerate=20
cell growth.}}

=20

= Fig. 1. =20 shRNA-mediated knockdown of tenascin-C does not = affect cell=20 proliferation in LN229 glioblastoma cells. (a) Lentiviral = shRNA=20 against control and tenascin-C was introduced into LN229 = cells and=20 IRES-mediated EGFP expressions were seen as an infection = marker.=20 Fluorescence (EGFP) and phase-contrast (PH) images are = shown. (b)=20 Representative immunoblot of tenascin-C and α-tubulin = in=20 shRNA-mediated protein knockdown against control and = tenascin-C in=20 LN229 cells. (c) Quantification of cell growth in control = and=20 tenascin-C knockdown cells 2 and 4 days after cell = plating.=20 Data are shown as mean =B1 SD (n =3D 3). Scale bar, = 200 =B5m; TNC-KD,=20 tenascin-C-knockdown cells.

[Normal=20 View ]

=20

Endogenous tenascin-C = regulates cell=20 motility in LN229 glioblastoma cells. Addition of = extrinsic=20 tenascin-C promotes glioma cell migration and=20 invasion.⁽¹⁸^{)=20
We tested whether endogenous tenascin-C expression is similarly =
involved=20
in glioma cell motility, and found that tenascin-C knockdown =
inhibited=20
glioma cell migration in the monolayer wound healing assay (Fig. 2a,b).=20
Cell migration requires remodeling of the actin cytoskeleton and =
assembly=20
and disassembly of focal adhesions.⁽¹²^{)=20
For this reason, we hypothesized that endogenous tenascin-C =
knockdown=20
alters cell adhesion, resulting in diminished cell motility. We =
examined=20
the cytoskeletons induced by endogenous tenascin-C knockdown. On =
the=20
fibronectin, the formation of focal adhesion is decreased in the =
control=20
cells, but it was not clearly changed in tenascin-C-knockdown =
cells, which=20
is consistent with that of past studies. Interestingly, tyrosine=20
phosphorylation of focal adhesion is decreased in =
tenascin-C-knockdown=20
cells on non-coated coverslips (Fig. 2c=96e).=20
It has been reported that tenascin-C suppresses RhoA activity and =
inhibits=20
the formation of actin stress fibers.⁽³⁰^{)=20
Therefore, we examined the involvement of RhoA in this effect on =
cell=20
migration; however, no apparent difference was detected in =
GTP-RhoA=20
(active form of RhoA) (data not shown). In focal adhesions, FAK =
and=20
p130 Cas (Crk-associated substrate) colocalize with=20
integrins.⁽¹³^{)=20
We found that tyrosine phosphorylation of FAK, but not =
p130 Cas, was=20
decreased in tenascin-C-knockdown cells. When we analyzed the =
LN229 cell=20
locomotion by cell-tracking analysis, tenascin-C knockdown glioma =
cells=20
significantly decreased the mean velocity and the distance from =
the=20
origin. Another question was whether or not cytoplasmic tenascin-C =
can act=20
directly on cell motility. Strikingly, coating the tissue culture =
dishes=20
with tenascin-C restored the migration velocity of the tenascin-C=20
knockdown cells to the level of the parent cells (Fig. 3),=20
which indicates that tenascin-C regulates glioblastoma cell =
migration in=20
an autocrine manner.}}}}

=20

= Fig. 2. =20 Endogenous tenascin-C regulates cell motility in = LN229=20 glioblastoma cells. (a) The confluent monolayers of the = control and=20 tenascin-C-knockdown LN229 cells were photographed 0 and = 14 h=20 after scraping. (b) The distance of cell-edge movements = 14 h=20 after scraping is shown as mean =B1 SD (n =3D 3). (c) F-actin = (phalloidin) and=20 tyrosine-phosphorylated proteins were stained in the control = and=20 tenascin-C-knockdown cells on the non-coated and fibronectin = coated=20 coverslips. (d,e) The average area and intensity of=20 tyrosine-phosphorylation signals in the control and = tenascin-C=20 knockdown cells stained by phosphor-tyrosine antibody on the = non-coated glass-bottom dishes are measured and shown as=20 mean =B1 SE (control, n =3D 30;=20 TNC-KD, n =3D 36). (f) = Representative=20 immunoblot of phosphorylated p130Cas and FAK in = shRNA-mediated=20 protein knockdown against control and tenascin-C in LN229 = cells.=20 TNC-KD, tenascin-C-knockdown cells; **, P < 0.001; scale bar,=20 200 =B5m.

[Normal View=20 ]

=20

= Fig. 3. =20 Endogenous tenascin-C activates glioblastoma cell = locomotion.=20 Cell tracking of (a) control cells, (b) tenascin-C-knockdown = cells=20 and (c) tenascin-C-knockdown with addition of exogenous = tenascin-C.=20 Cells were imaged every 5 min for 12 h under = microscopy.=20 (d) Track length (=B5m). (e) Mean velocity (=B5m/min). (f) = Distance from=20 the origin. Tracks and velocities were calculated with = MetMorph=20 software (Molecular Devices, Sunnyvale, CA). TNC-KD,=20 tenascin-C-knockdown cells; TNC-KD + TNC,=20 tenascin-C-knockdown cells with exogenous tenascin-C; *, = P < 0.01; **, P < 0.001; scale bars,=20 50 =B5m.

[Normal View=20 ]

=20

Tenascin-C-knockdown = decreases=20 glioblastoma cell invasiveness and reactive change of peritumoral = brain=20 tissue. We next examined whether endogenous tenascin-C = is=20 involved in glioblastoma cell invasion in = vivo. We=20 used a xenograft model implanting control and tenascin-C-knockdown = LN229=20 cells into the striatum of BALB/c nude mice. Using this model, we=20 evaluated the invasiveness of glioblastoma cells 3 weeks = after=20 implantation. Control shRNA cells exhibited tumor growth and = infiltration=20 into the surrounding brain tissue, forming small tumor clusters = distinct=20 from the tumor core (Fig. 4a,c).=20 Histopathologically, edematous change was observed in the brain = parenchyma=20 containing glioblastoma infiltration (Fig. 4b).=20 Intercellular tenascin-C was also diffusely deposited not only in = the=20 tumor core but also in the surrounding brain tissue with = glioblastoma cell=20 infiltration (Fig. 4d).=20 In contrast, although tumor size of tenascin-C-knockdown LN229 = cell was=20 approximately equal to that of control cells (Fig. 4i),=20 the number of infiltrating glioma cells and tumor clusters in the=20 surrounding brain were decreased (Fig. 4g).=20 As expected, these cells deposited significantly less tenascin-C = than the=20 control (Fig. 4h),=20 and the edematous change was decreased in the brain parenchyma = around the=20 tumor (Fig. 4f).=20 The infiltrating control cells show dense cytoplasmic tenascin-C = (Fig. 4c,d,=20 white rectangles). Quantification of tumor cluster number revealed = that=20 tenascin-C knockdown decreased cluster formation by approximately = half,=20 compared with control cells (Fig. 4j).=20 These results indicate that endogenous tenascin-C promotes = glioblastoma=20 cell invasion and the adjacent reactive change of the brain = parenchyma but=20 is not involved in tumor growth in the glioblastoma xenograft=20 models.

=20

= Fig. 4. =20 Endogenous tenascin-C enhances glioblastoma invasion = with=20 reactive change of the brain tissue in a xenograft = glioblastoma=20 model. Intracranial tumors derived from (a,b,c,d) control = cells and=20 (e,f,g,h) tenascin-C-knock down cells are shown by = hematoxylin=96eosin=20 staining (HE), IRES-mediated EGFP expression of implanted = cells=20 (EGFP) and tenascin-C staining (TNC). Small and large = rectangles in=20 (a) and (e) are enlarged in (b,f) and (c,d,g,h), = respectively. Small=20 white rectangles are enlarged to large white rectangles in = (c,d).=20 (i) Tumor size is shown as mean =B1 SD (n =3D 6). (j) Number of cell = clusters per=20 circumference of the tumor core is shown as = mean =B1 SD=20 (n =3D 6). Scale bars, = 1 mm (a,e),=20 100 =B5m (b,f), 500 =B5m (c,d,g,h); *, P < 0.01. =

[Normal=20 View ]

=20

Tenascin-C expression = correlates with=20 the volume of the reactive brain tissue in = glioblastoma. Our=20 experimental study showed that endogenous tenascin-C promotes = glioblastoma=20 invasion and a secondary reactive change of the peritumoral = tissue. The=20 reactive change of brain tissue contains vasogenic edema and = compositional=20 change of brain tissue induced by tumor cell infiltration. In = modern MR=20 imaging, the tumor core is enhanced by gadolinium and the = peritumoral=20 change in brain parenchyma is seen as a hyperintense region by = FLAIR=20 sequences (Fig. 5a).=20 It is therefore possible that the peritumoral hyperintensity area=20 represents the secondary change induced by tenascin-C-facilitated=20 glioblastoma cell invasion. We examined the correlation between = the=20 FLAIR/Enhanced volume (F/E) ratio (indicating the volume of = peritumoral=20 tissue change), with tenascin-C mRNA expression in resected tumors = from 16=20 glioblastoma patients. Strikingly, tenascin-C expression = significantly=20 correlates with the volume of the peritumoral tissue change = (Spearman=20 correlation test, P =3D 0.005, = r =3D 0.77) (Fig. 5b),=20 although it does not correlate with the proliferation index by a = mitotic=20 marker Ki-67 staining (Spearman correlation test, P =3D 0.272, r =3D =960.29) (Fig. 5c).

=20

= Fig. 5. =20 Correlation between tenascin-C expression and = reactive change=20 of peritumoral brain parenchyma in resected glioblastomas. = (a)=20 Representative MR images of glioblastoma show the tumor core = enhanced by gadolinium (Gd enhanced) and the peritumoral = change of=20 brain parenchyma as hyperintense area of FLAIR sequence. (b) = Correlation between FLAIR/Enhanced volume ratio and = tenascin-C mRNA=20 expression in 16 patients with glioblastoma. (c) Correlation = between=20 proliferation activity (Ki-67 index) and tenascin-C = expression in 16=20 resected glioblastomas. a. u., arbitrary unit; r, Spearman correlation coefficient. =

[Normal View=20 ]

The localization of tenascin-C in resected = glioblastomas has=20 been described in the perivascular and intercellular spaces. = However, it=20 is unclear to what extent tenascin-C is expressed around the=20 tumor.⁽³¹^{)=20
We therefore examined the localization of tenascin-C in resected=20
glioblastoma sections obtained from either the tumor core=20
(gadolinium-enhanced area) or the surrounding lesion =
(hyperintensity area=20
of FLAIR imaging) by using intraoperative navigation systems. In =
patients=20
with high and low F/E ratios, intercellular deposition of =
tenascin-C was=20
detected in the tumor core (Fig. 6a,b,=20
upper panels, TNC). The peritumoral brain parenchyma, showing=20
hyperintensity with FLAIR imaging, contained glioblastoma cells =
undergoing=20
mitosis (Fig. 6a,b,=20
lower panels, Ki-67). In the patient with a high F/E ratio, a =
number of=20
glioblastoma cells infiltrated in the peritumoral brain tissue and =

exhibited high deposition of tenascin-C in their cytoplasm (Fig. 6b,=20
lower panels). Based on these results, one possible mechanism of=20
glioblastoma progression is that glioblastoma infiltration into =
the brain=20
parenchyma is enhanced by overexpression of endogenous tenascin-C, =
which=20
induces reactive changes of the brain tissues, detected as =
hyperintensity=20
area with FLAIR sequences.}

=20

= Fig. 6. =20 Infiltrating glioblastoma cells show cytoplasmic = deposition=20 of tenascin-C in reactive changes of peritumoral brain = parenchyma=20 detected by FLAIR image. Representative images of the tumor = core and=20 peritumoral tissue in patients with (a) low FLAIR/Enhanced = volume=20 ratio (F/E ratio) and (b) high F/E ratio. Each square in MR = images=20 (left panel) indicates the localization of the microscopic = images=20 (three right panels; HE, Ki-67, TNC). Hematoxylin=96eosin = staining=20 (HE), mitotic activity staining (Ki-67), tenascin-C staining = (TNC).=20 Scale bars, 200 =B5m.

[Normal=20 View = ]

=20

=20

Discussion

Numerous studies have revealed that tenascin-C = plays a=20 certain role in tumor progression including tumorigenesis, = proliferation,=20 invasion, metastasis and angiogenesis.^{(14,15,18,19,22,32=9638)}=20 It is widely acknowledged that tenascin-C disrupts the binding of=20 fibronectin to the integrin=20 = α₅β₁/syndecan-4= complex at=20 the cell surface, which causes morphological change of cultured = cells and=20 promotes cell proliferation.⁽^22,39,40⁾=20 However, in this study, no significant differences exist in cell=20 morphology or proliferation between the control and = tenascin-C-knockdown=20 glioblastoma cells. This is possibly because tenascin-C affects = cell=20 growth and morphology in an environment containing extrinsic = tenascin-C=20 and fibronectin.(^20,33^{)<=
/SUP>=20
While collagen, fibronectin and laminin are the predominant ECM =
components=20
in non-central nervous system tissue, major ECM components of =
brain=20
parenchyma include hyaluronan, proteoglycans and =
glycosaminoglycans, such=20
as hyaluronic acid and chondroitin sulfate. Fibronectin is found =
at the=20
gliomesenchymal junction of tumors and in tumor-associated blood=20
vessels.⁽⁴¹^{)=20
Glioblastomas do not express fibronectin, yet fibronectin is =
confined to=20
proliferating vessel walls and =
leptomeninges.⁽⁴²^{)=20
Fibronectin was either localized in the basement membrane or =
formed thick,=20
multi-layered deposits in the vessel walls. Fibronectin expression =
was=20
seen in all glioblastoma samples but its distribution was not =
uniform, and=20
its staining intensity was weak.⁽³¹^{)=20
These observations suggest that endogenous tenascin-C secreted by=20
glioblastoma cells does not affect cell morphology and =
proliferation when=20
the tumor microenvironment does not abound in fibronectin.}}}}

In this study, glioblastoma cell motility on a = non-coated=20 substrate was significantly suppressed in tenascin-C-knockdown = cells,=20 indicating that endogenous tenascin-C accelerates glioblastoma = cell=20 invasion via different machinery from the action of tenascin-C on=20 fibronectin. The same phenotype was also seen in a xenograft model = whereby=20 tenascin-C-knockdown glioblastoma cells exhibited less = invasiveness into=20 brain parenchyma. Immunohistochemical localization of tenascin-C = in vivo experiments and clinical samples = demonstrate=20 predominant cytoplasmic deposition in the glioblastoma cells = invading into=20 brain tissue, especially in the patient with high F/E ratio, which = shows=20 highly invasive tumor. These observations suggest that = glioblastoma cells=20 with abundant expression of tenascin-C invade aggressively and = secreted=20 tenascin-C acts to enhance their invasiveness in an autocrine = manner.=20 Tenascin-C was described as a hexabrachion because it exists as a = hexamer=20 of 220=96320-kDa subunits. Each subunit of tenascin-C comprises = three types=20 of structural modules: EGF-like domains, fibronectin type III = (FNIII)=20 repeat domains, and a terminal knob homologous to the beta- and=20 gamma-chains of fibrinogen. This multidomain structure of the = tenascin-C=20 suggests the possibility of multiple independent=20 functions.⁽⁴³^{)=20
In this study, we confirmed that extrinsic tenascin-C rescued the=20
migration disorder of tenascin-C knockdown cells. It is a possible =

mechanism that endogenous tenascin-C is involved in the regulation =
of=20
focal adhesion turnover.}

The expression level of tenascin-C correlates with = prognosis=20 in patients with many malignancies.^{(14,37,44=9649)}=20 In the previous study, we identified tenascin-C expression as = prognosis=20 factor of patients with glioblastoma.⁽⁹⁾= =20 The invasive nature of gliomas also causes breakdown of the = blood=96brain=20 barrier and cerebral edema formation, which is responsible for = significant=20 morbidity and mortality in patients with glioblastoma. In this = study, we=20 found that the expression level of tenascin-C correlates with the = degree=20 of peritumoral change in brain parenchyma. This change in brain = parenchyma=20 is thought to be comprised of focal brain edema and changes of=20 extracellular matrix following glioblastoma cell invasion; = however, the=20 underlying mechanism is still unclear.

The change of the peritumoral brain tissue is = usually=20 detected as hyperintensity in T2-weighted images and FLAIR = sequences of MR=20 imaging.⁽^2,3,7^{)=20
Regrettably, it was impossible to evaluate discriminatively the=20
gadolinium-enhanced tumor core and the peritumoral hyperintensity =
area in=20
our mouse xenograft model. In the immunohistochemical analysis, =
diffuse=20
deposition of tenascin-C was observed in the stroma of the tumor =
core and=20
peritumoral brain tissue. Interestingly, strong cytoplasmic =
deposition of=20
tenascin-C was detected in the migrating glioblastoma cells at the =

invasion border and in the peritumoral brain tissue. In a study of =
breast=20
cancer cells, Ishihara et al. emphasized that cancer spreading might be =
involved in=20
cytoplasmic deposition of tenascin-C, but not in intercellular=20
deposition.⁽⁵⁰^{)=20
Taken together, tenascin-C may be one of the key extracellular =
matrix=20
components facilitating glioblastoma cell invasion and promoting =
the=20
peritumoral change of brain parenchyma.}}

In summary, we report that endogenous tenascin-C = facilitates=20 glioblastoma cell invasion by regulating focal adhesion and that=20 glioblastoma cells with high tenascin-C expression exhibit = invasive=20 nature. Consistent with these observations, the expression level = of=20 tenascin-C correlates with the degree of peritumoral tissue = change,=20 detected as FLAIR hyperintensity in clinical samples. The detail = molecular=20 basis for this curious ability of endogenous tenascin-C to = facilitate=20 glioblastoma cell invasion is still unclear and warrants further=20 investigation. This study highlights tenascin-C's attractiveness = as a=20 target molecule for anti-invasion therapy in the treatment of=20 glioblastoma.

=20

=20

=20

Acknowledgments

We thank M Way (Cancer Research UK, London) for = reagents. We=20 thank members of Arakawa's and Matsuda's laboratories for = constructive=20 comments on the manuscript and throughout the project. This work = was=20 supported by grants from a Grant-in-Aid for Specially Promoted = Research=20 from the Ministry for Education, Culture, Sports, Science and = Technology=20 of Japan (YA, YK, JAT, MM, NH), Kyoto University Research Fund = (YA) and=20 Japan Heart Foundation/Bayer Yakuhin Research Grant = (YA).

=20

=20

Disclosure Statement

We declare that we have no competing financial=20 interests.

=20

=20

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