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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_01C9F748.B913D340 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_01C9F748.B913D340 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
Matherials and Methods
Results
Discussion
Acknowledgments
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. Resveratrol (Rsv) = and quercetin (Quer) reduced the number of human, mouse, and rat = gliomas. Glioma cell lines from rat (C6), mouse (GL261), and humans = (U87-MG and U138-MG) were treated with 10 or 50 µM Rsv, 25 or = 100 µM Quer, and 10 µM Rsv plus 25 µM = Quer for 24, 48, and 72 h; cells were then counted in a = hemocytometer. Each time point represents the = mean ± SEM of three independent experiments. anova was = followed by SNK post test. *P < 0.05, = **P < 0.01, ***P < 0.001.'); }=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. Resveratrol (Rsv) = increased propidium iodide (PI) incorporation in C6 cells but not in = astrocyte cell cultures. C6 cells were treated with 10 or = 50 µM Rsv, 25 or 100 µM quercetin (Quer), and = 10 µM Rsv plus 25 µM Quer. Astrocyte cultures were = treated with 1, 10, 25, or 50 µM Rsv. At the end of the = treatments (72 h), the images were obtained using a Carl Zeiss = inverted microscope. Representative light micrographs of C6 (a), = astrocyte cell cultures (c), fluorescence micrographs of C6 (b), and = astrocyte cell cultures (d) stained with PI (×100) are shown. The = numbers below the pictures depict values of PI staining (ratio of PI = labeled cells/total number cells, normalized to control) as = mean ± SEM of three independent experiments. anova was = followed by SNK post test ***P < 0.001.'); }=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. Resveratrol (R) = induces caspase 3/7 activation, annexin V staining, and redudes mitotic = index. C6 cells were treated with Rsv (10, 25, or 50 µM) and = quercetin (Q) (25 or 100 µM) and 10 µM Rsv plus = 25 µM Quer for 48 or 72 h. Caspase 3/7 activity was = estimated by fluorogenic assay for the effector caspase 3/7 (a). Cells = were stained with annexin V-FLUOS and propidium iodide (PI) and positive = cells were observed using an inverted fluorescence microscope = (×200). The same fields are shown. Control cells (left panels), = 50 µM Rsv (middle panels), and 10 µM Rsv plus = 25 µM Quer (right panels) observed under light microscopic = image (upper panel), with annexin V staining (middle panel) and PI = staining (lower panel) after 72 h of treatments (b). Cells were = treated as indicated above for 24, 48, and 72 h and cells in = mitosis were visualized by PI staining. The results represent = mean ± SEM from three independent experiments. anova = was followed by SNK post test. *P < 0.05, = **P < 0.01, ***P < 0.001.'); }=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. Resveratrol = (Rsv) and quercetin (Quer) decreased the phosphorylation of Akt. C6 = cells were treated with 10 µM Rsv, 25 µM Quer, or = 10 µM Rsv plus 25 µM Quer, for 30 min, = 4 h, and 24 h, and with 10 µM LY294002 for = 30 min. Lysates were subjected to western blot analysis for = pS473Akt and total Akt antibodies (a) and PTEN antibody (b). = Representative blots of three independent experiments are shown and the = numbers below the blots depict values of densitometric evaluation as = mean ± SEM. Number of C6 cells treated as indicated for = 72 h (c). The results represent mean ± SEM from = three independent experiments. anova was followed by SNK post test. = *P < 0.05, **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. Resveratrol (Rsv) = and quercetin (Quer) induce senescence in C6 cells. C6 cells were = cultivated in the continuous presence of 10 or 25 µM Rsv, = 25 µM Quer, 10 µM Rsv plus 25 µM Quer, or 10 = µM LY294002 for 12 days and cumulative population doublings were = plotted against time (a). After this time cells were stained for = senescence-associated-β-galactosidade (SA-β-gal) and = photographed (b). The bar graph represents the ratio of = SA-β-gal-positive cells to total cells of at least three fields = of three independent experiments (c). Colony formation efficiency of C6 = cells 7 days after 12 days Rsv or Quer treatment or 6 days of = cotreatment (d), and SA-β-gal staining at the end of the colony = formation assay (e). anova was followed by SNK post test. = *P < 0.05, **P < 0.01, = ***P < 0.001.'); }=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= =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('');hwnd.document.write('

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3D""=20

Resveratrol and quercetin cooperate to = induce=20 senescence-like growth arrest in C6 rat glioma cells

Lauren=20 L. Zamin = ¹=20 , Eduardo = C. Filippi-Chiela ¹ = ,=20 Patricia Dillenburg-Pilla ³ = ,=20 Fabiana Horn ¹ , = Christianne Salbego ² = and Guido Lenz ^1,4

¹ Biophysics = Department=20 , = ² Biochemistry = Department,=20 Universidade Federal do Rio Grande do Sul (UFRGS), Porto = Alegre ;=20 = ³ Biochemistry = Department,=20 Universidade de S=E3o Paulo (USP), Ribeirao Preto, Brazil=20

⁴ To whom = correspondence=20 should be addressed. E-mail: lenz@ufrgs.br =

Copyright =A9 2009 Japanese Cancer=20 Association

ABSTRACT

3DAbstract=20

Glioma is the most frequent and malignant primary = human=20 brain tumor with dismal prognosis despite multimodal therapy. = Resveratrol=20 and quercetin, two structurally related and naturally occurring=20 polyphenols, are proposed to have anticancer effects. We report = here that=20 resveratrol and quercetin decreased the cell number in four glioma = cell=20 lines but not in rat astrocytes. Low doses of resveratrol = (10 =B5M) or=20 quercetin (25 =B5M) separately had no effect on apoptosis = induction,=20 but had a strong effect on caspase 3/7 activation when = administered=20 together. Western blot analyses showed that resveratrol = (10 =B5M) and=20 quercetin (25 =B5M) caused a reduction in phosphorylation of = Akt, but=20 this reduction was not sufficient by itself to mediate the effects = of=20 these polyphenols. Most important, resveratrol and quercetin = chronically=20 administered presented a strong synergism in inducing = senescence-like=20 growth arrest. These results suggest that the combination of = polyphenols=20 can potentialize their antitumoral activity, thereby reducing the=20 therapeutic concentration needed for glioma treatment. (Cancer Sci 2009)

(Received March 26, 2009/Revised April 30, = 2009/Accepted May=20 7, 2009)

DIGITAL OBJECT IDENTIFIER (DOI)

10.1111/j.1349-7006.2009.01215.x About=20 DOI

=20

=20

Article Text

Malignant gliomas comprise the most common and = devastating=20 types of primary central nervous system (CNS) tumors and have an = incidence=20 of 5=968/100 000 population. The median survival of patients = with=20 malignant gliomas is only 12 months after=20 diagnosis.⁽¹⁾= =20 Gliomas are characterized by diffuse infiltrations of the = surrounding CNS=20 tissue,⁽²⁾= =20 and this characteristic is one of the main reasons for the lack of = successful treatment of this type of = cancer.⁽³⁾= =20 Nevertheless, the therapeutic strategy for gliomas has remained=20 essentially unchanged for decades due to a limited understanding = of the=20 biology of the disease. There is an urgent need to develop = mechanism-based=20 approaches for the management of glioma, i.e. to develop = strategies which=20 can eliminate only the damaged (pre-malignant) or malignant cells = without=20 harming normal CNS tissue.⁽⁴⁾=

Resveratrol (Rsv; = 3,5,4'-trihydroxy-trans-stilbene) is found=20 in many plant species such as grapes, berries, and peanuts, and = exhibits=20 pleiotropic health beneficial effects, including antioxidant,=20 anti-inflammatory, cardioprotective, neuroprotective, and = antitumor=20 activities.⁽⁵⁼⁹⁶⁷^{)=20
The cancer chemopreventive properties of Rsv were first =
appreciated when=20
Jang et al.=20
(1997)⁽⁵⁾=
=20
demonstrated that Rsv inhibited cellular events associated with =
tumor=20
initiation, promotion, and progression. It has been shown that Rsv =
can=20
induce apoptotic cell death in a number of cancer cell lines, =
including=20
U251 human glioma cells,⁽⁸⁾=
=20
RT-2 rat glioma cells,⁽⁹⁾=
=20
and the C6 rat glioma cell line.⁽¹⁰^{)=20
It is thought that Rsv affects whole pathways and sets of =
intracellular=20
events rather than a single enzyme and is, therefore, less prone =
to induce=20
resistance in cancer cells.⁽¹¹⁾}}

Quercetin (Quer; 3,3',4',5,7-pentahydroxyflavone) = is a=20 flavonoid ubiquitous in nature that has recently been described as = a=20 potential anticancer agent⁽¹²^{)=20
because of its ability to modulate cell=20
proliferation,⁽¹³^{)=20
survival, and differentiation, targeting key molecules responsible =
for=20
tumor cell growth such as p53, p21, and Ras in several cancer cell =

types.⁽^14,15^{)<=
/SUP>=20
Quer induces little or no toxicity when administered orally or=20
intravenously, and preliminary animal and human data indicate that =
Quer=20
inhibits tumor growth.⁽¹⁶^{)=20
We have previously demonstrated that Quer induced necrotic and =
apoptotic=20
cell death in the U138-MG human glioma cell=20
line,⁽¹⁷^{)=20
and Kim et al. =
(2007)=20
demonstrated that Quer treatment resulted in loss of cell =
viability in the=20
A172 glioma cell line.⁽¹⁸⁾}}}}}

Fresh grape skin contains about = 50=96100 =B5g/g of Rsv and=20 40 =B5g/g of Quer.⁽^19,20^{)<=
/SUP>=20
In red wine, the concentration of Rsv is in the range of=20
7=9613 =B5M⁽²¹^{)=20
and Quer is in the range of =
7.4 =B5M,⁽²²^{)=20
while commercial grape juice contains about 20 =B5M of trans-resveratrol.⁽²³^{)=20
In rats, injection of Rsv or Quer lead to levels of these drugs in =
the low=20
micromolar range in the CNS, suggesting that they could =
potentially be=20
used as therapeutic agents against =
gliomas.⁽^24,25^)<=
/SUP>}}}}

The mechanisms of chemopreventive effects of Rsv = and Quer=20 are not well understood. Moreover, studies suggest that targeting=20 regulatory molecules of the cell cycle, apoptosis, and senescence = in=20 cancer cells can be an effective strategy for the management of=20 cancer⁽^26=9628⁾=20 and, at present, many works have shown that Rsv and Quer can = modulate some=20 of these processes.(^17,29=9631⁾=20 Both Rsv and Quer have been suggested to extend lifespam of model=20 organisms(^32=9635⁾=20 and to reduce cell senescence in normal = cells.(^36,37^{)<=
/SUP>=20
There is only one report showing that Rsv can induce senescence in =
cancer=20
cells⁽³¹^{)=20
and no such report was found for Quer. Besides, no study =
considering the=20
interactions of these agents on cell senescence was found in the=20
literature. Here we show that Rsv and Quer cooperate in triggering =

apoptosis, inhibiting cell proliferation and, most strikingly, in =
inducing=20
senescence-like growth arrest in gliomas cell lines.}}

=20

=20

Matherials and Methods

=20

Cell = culture. Rat C6=20 glioma cells, human U87-MG, and U138-MG glioma cells were obtained = from=20 ATCC (Rockville, MD, USA). Mouse GL261 glioma cell line was gently = provided by Dr Ilker Ey=FCpoglu, University of Zurich. All culture = materials=20 were purchased from Gibco Laboratories (Grand Island, NY, USA). = Cells were=20 cultured in DMEM low glucose supplemented with 10% FBS (5% for C6 = cells),=20 1% penicillin/streptomycin, and 0.1% anphoterecin B at 37=B0C/5%=20 CO₂ in a humidified incubator.

=20

Cell = treatments. Rsv and=20 Quer were purchased from Sigma Chemical (St. Louis, MO, USA) and=20 administered for 24=9672 h. Rsv (10 =B5M) and Quer = (25 =B5M) were=20 also added together and are referred in the text as cotreatment. = The PI3K=20 (Phosphoinositide3-kinase) inhibitor LY294002 (LY; Cell Signaling, = Beverly, MA, USA), was administered (10 =B5M) for 72 h. = The=20 vehicle carrier DMSO (Acros Organics, NJ, USA) was present in the = control=20 wells at a maximum concentration of 0.5%.

=20

Astrocyte=20 cultures. Primary astrocyte cultures were prepared as=20 previously described.⁽³⁸^{)=20
Briefly, hippocampi of new-born Wistar rats (1=962 days old) were =
removed=20
and mechanically dissociated by sequential passage through a =
Pasteur=20
pipette. Cell suspension was centrifuged at 1000 g for 5 min. The supernatant was =
discarded,=20
and the pellet resuspended in culture medium containing DMEM with =
10% FBS=20
and 1 % penicillin/streptomycin and 0.1% anphoterecin B. Cells =
were plated=20
at a density of 1.5 =D7 10⁵=20
cells/cm² onto 24-well plates pretreated =
with=20
poly-lysine in 5% CO₂ at 37=B0C, and =
allowed to grow=20
to confluence (14 days). All animal use procedures were approved =
by the=20
local Animal Care Committee and were in accordance with the NIH =
Guide for=20
the Care and Use of Laboratory Animals.}

=20

Cell number determination = and=20 propidium iodide (PI) staining. The number of cells = was=20 counted in a hemocytometer and the value of the treated cells was=20 calculated as a percentage of the control. For PI staining, cells = were=20 treated with the drugs in the presence of 6 =B5M PI (Sigma) = for=20 72 h and images were obtained using a Carl Zeiss Fluorescence = Inverted Microscope with a rhodamine filter (Carl Zeiss, Jena, = Germany).=20 Results are presented as ratio of cells marked with PI to total = cells of=20 at least three fields of three experiments analyzed by Image J = software=20 (NIH Image, Rockville, MD, USA).

=20

Mitotic index (MI)=20 determination. Cells were fixed with 70% ethanol for=20 15 min and stained with 6 =B5M PI. MI was evaluated by = counting at=20 least 300 cells per sample using Image J software and the = percentage of=20 cells with condensed chromatin was determined.

=20

Long-term culture of C6 = cells in the=20 presence of Rsv and Quer and determination of population=20 doublings. Cells were plated at each passage in = 24-well plates=20 at a density of 50 000 cells/well and exposed to DMSO or = different=20 concentrations of Rsv and Quer with renewal every 48 h of = medium and=20 treatments. Confluent cells were passaged, and population = doublings (PD)=20 were determined according to the formula PD =3D[log N(t)*logN(to)]/log 2, where N(t) is the number of cells per well at time of = passage, and=20 N(to) is the number = of cells=20 seeded at the previous passage.⁽³¹^{)=20
The sum of PDs was then plotted against time of =
culture.}

=20

Colony formation=20 assay. After 6 or 12 days of treatments, cells were=20 trypsinized, counted, and plated at concentration of 1000 = cells/well into=20 six-well plates. After 7 days without treatments, senescence = induction was=20 analyzed as described below and cells were stained with Giemsa = solution to=20 determinate colony number formation.⁽³⁹^{)=20
The percentage of colony forming efficiency was calculated in =
relation to=20
values of untreated control cells.}

=20

Senescence-associated-β-galactosidade=20 (SA-β-gal) staining. After 12 days in culture, = cells were=20 washed in PBS, fixed in 3% formaldehyde for 15 min at room=20 temperature, washed, and incubated with fresh SA-β-gal = staining solution=20 containing 1 mg/mL X-gal (Sigma), 40 mM citric = acid/sodium=20 phosphate (pH 6.0), 5 mM potassium ferrocyanide, = 5 mM=20 potassium ferricyanide, 150 mM NaCl, and 2 mM MgCl for=20 12=9616 h at 37=B0C.⁽⁴⁰^{)=20
Results are presented as ratio of SA-β-gal-positive cells to =
total cells=20
of at least three fields of three experiments.}

=20

Caspase 3/7=20 activity. Measurement of caspase 3/7 activity was done = as=20 previously described.⁽⁴¹^{)=20
After treatments, C6 cells were lysed and 40 =B5g of protein =
were=20
incubated with a reaction buffer. The caspase 3/7 substrate =
Ac-DEVD-MCA=20
(Peptide Institute, Osaka, Japan) was present at a final =
concentration of=20
20 =B5M. The cleavage of the fluorogenic synthetic substrate =
was=20
quantified by fluorimetry in a Spectramax Microplate Reader =
(Molecular=20
Devices, Sunnyvale, CA, USA). The fluorescence intensity was =
calibrated=20
with standard concentrations of MCA, and the caspase 3/7 activity =
was=20
calculated using the slope of the accumulated fluorogenic product =
and=20
expressed in picomols per min per mg of protein. Cells treated =
with=20
50 =B5M etoposide (Sigma) for 24 h were used as positive =

control.}

=20

Annexin V=20 staining. Phosphatidylserine externalization was = determined by=20 annexin fluorescent signal of Annexin-V-FLUOS Staining Kit assay = (Roche,=20 Germany) according to the manufacturer's protocol. C6 cells were = treated=20 and incubated with PI and annexin for 10 min at room = temperature and=20 photographed. Necrotic cells were identified by double PI and = annexin=20 fluorescent staining, while apoptotic cells were positive only for = annexin.

=20

Immunoblotting. Preparation of = protein samples=20 and western blot were performed as described previously with minor = modifications.⁽⁷⁾= =20 Whole-cell lysate (40 =B5g) was electrophoresed and = electroblotted onto=20 a PVDF membrane (Amersham Pharmacia Biotech, Piscataway, NJ, USA). = The=20 membrane was blocked in 5% skimmed milk/TBST for an hour and = probed with=20 Ab to phosphatase and tensin homolog (PTEN) (1:2000 dilution; Cell = Signaling), phospho-Akt (pAkt Ser473) (1:1000; Cell Signaling), = and Akt=20 (1:1000; Cell Signaling) for 1 h at room temperature. Bound = Ab was=20 detected with appropriate HRP secondary Ab (1:2000; Cell = Signaling) using=20 ECL and X-ray films (Kodak X-Omat, Rochester, NY, USA). The = optical=20 density of the bands was obtained in with Bio-Rad software = (Quantity One;=20 Hercules, CA).

=20

Statistical = analysis. All=20 data were expressed as mean =B1 SEM and were analyzed = using anova, followed by SNK post-hoc test = for=20 multiple comparisons. Differences with P < 0.05 were considered = significant. Analyses=20 of the data were performed using the software GraphPad INSTAT, = (GraphPad=20 Software, San Diego, CA, USA).

=20

=20

Results

=20

Rsv and Quer decreased = cell=20 numbers. In search of novel strategies to induce cell = death,=20 we investigated the antitumor effect of the natural compounds Rsv = and Quer=20 on glioma cell lines. As shown in Figure 1,=20 Rsv and Quer decreased the number of glioma cells in a time- and=20 dose-dependent manner. In C6 cells, after 72 h of = 10 =B5M Rsv and=20 25 =B5M Quer treatment, the number of cells was reduced 25% = and 54%,=20 respectively. When cells were treated with both compounds at the = same=20 concentration and exposure time, the number of cells was reduced = 80%,=20 indicating an additive effect of the cotreatment. This reduction = was=20 similar to the result observed with higher concentrations of Rsv = and Quer.=20 These results were confirmed by MTT assay (data not shown). In = other=20 glioma cell lines, the additive effect was not observed, with = 50 =B5M=20 Rsv being the dose that induced the biggest cell number reduction = after=20 72 h of treatment. The difference of response could be = explained by=20 the genetic characteristics of each cell line, since U87-MG and = U138-MG=20 are deleted for PTEN, and U138-MG is mutant for TP53, whereas C6 = has both=20 proteins functional.⁽^42,43^)<=
/SUP>

=20

= Fig. 1. =20 Resveratrol (Rsv) and quercetin (Quer) reduced the = number of=20 human, mouse, and rat gliomas. Glioma cell lines from rat = (C6),=20 mouse (GL261), and humans (U87-MG and U138-MG) were treated = with 10=20 or 50 =B5M Rsv, 25 or 100 =B5M Quer, and = 10 =B5M Rsv plus=20 25 =B5M Quer for 24, 48, and 72 h; cells were then = counted=20 in a hemocytometer. Each time point represents the=20 mean =B1 SEM of three independent experiments. = anova was followed by SNK post = test.=20 *P < 0.05, **P < 0.01, ***P < 0.001. =

[Normal=20 View ]

To evaluate whether the cytotoxic effect of Rsv is = selective=20 to glioma cell cultures, primary astrocyte cultures and glioma C6 = cells=20 were treated for 72 h with Rsv and stained with PI. As shown = in Figure 2(a,b),=20 PI incorporation was highest in glioma cells treated with = 50 =B5M Rsv.=20 This effect was not observed in rat astrocyte cultures (Fig. 2c,d).=20 C6 cells treated with 25=96100 =B5M Quer (Fig. 2a,b)=20 presented no significant PI incorporation.

=20

= Fig. 2. =20 Resveratrol (Rsv) increased propidium iodide (PI)=20 incorporation in C6 cells but not in astrocyte cell = cultures. C6=20 cells were treated with 10 or 50 =B5M Rsv, 25 or = 100 =B5M=20 quercetin (Quer), and 10 =B5M Rsv plus 25 =B5M = Quer. Astrocyte=20 cultures were treated with 1, 10, 25, or 50 =B5M Rsv. = At the end=20 of the treatments (72 h), the images were obtained = using a Carl=20 Zeiss inverted microscope. Representative light micrographs = of C6=20 (a), astrocyte cell cultures (c), fluorescence micrographs = of C6=20 (b), and astrocyte cell cultures (d) stained with PI = (=D7100) are=20 shown. The numbers below the pictures depict values of PI = staining=20 (ratio of PI labeled cells/total number cells, normalized to = control) as mean =B1 SEM of three independent = experiments.=20 anova was followed by SNK = post test=20 ***P < 0.001.=20

[Normal View=20 ]

=20

Rsv activated caspase=20 3/7. To assess whether cytotoxicity of Rsv and Quer = was due to=20 apoptosis induction, we measured the activity of the executioner = caspase=20 3/7⁽⁴⁴^{)=20
after treating C6 cells with these compounds. Rsv (50 =B5M) =
increased=20
caspase 3/7 activity after 48 h of treatment at an intensity =
similar=20
to 50 =B5M etoposide, used as a positive control (Fig. 3a).⁽⁴¹^{)=20
This activation was slow, since there was no detectable caspase =
3/7=20
activation before this time (data not shown). Quer (100 =B5M) =
was=20
unable to induce caspase 3/7 even at 72 h of treatment. =
Neither=20
10 =B5M Res nor 25 =B5M Quer alone were able to induce =
caspase 3/7=20
activation; however, a strong activation was observed with the=20
cotreatment, suggesting a synergistic effect of these compounds in =
caspase=20
3/7 activation (Fig. 3a).}}

=20

= Fig. 3. =20 Resveratrol (R) induces caspase 3/7 activation, = annexin V=20 staining, and redudes mitotic index. C6 cells were treated = with Rsv=20 (10, 25, or 50 =B5M) and quercetin (Q) (25 or = 100 =B5M) and=20 10 =B5M Rsv plus 25 =B5M Quer for 48 or 72 h. = Caspase 3/7=20 activity was estimated by fluorogenic assay for the effector = caspase=20 3/7 (a). Cells were stained with annexin V-FLUOS and = propidium=20 iodide (PI) and positive cells were observed using an = inverted=20 fluorescence microscope (=D7200). The same fields are shown. = Control=20 cells (left panels), 50 =B5M Rsv (middle panels), and = 10 =B5M=20 Rsv plus 25 =B5M Quer (right panels) observed under = light=20 microscopic image (upper panel), with annexin V staining = (middle=20 panel) and PI staining (lower panel) after 72 h of = treatments=20 (b). Cells were treated as indicated above for 24, 48, and = 72 h=20 and cells in mitosis were visualized by PI staining. The = results=20 represent mean =B1 SEM from three independent = experiments.=20 anova was followed by SNK = post=20 test. *P < 0.05, **P < 0.01, ***P < 0.001.

[Normal=20 View ]

To verify if the caspase 3/7 activation was = leading to=20 apoptosis, we analyzed annexin V (an apoptosis marker) and PI = (necrosis or=20 late apoptosis marker) staining. After 72 h of 50 =B5M = Rsv and=20 cotreatment of Rsv and Quer, there was an increase in annexin V = and PI=20 staining, suggesting that the observed caspase 3/7 activation was = leading=20 to apoptosis (Fig. 3b).

In order to investigate the effect of Rsv and Quer = on cell=20 cycle progression we determined the MI. The MI decreased with = 50 =B5M=20 Rsv and 100 =B5M Quer treatments but did not decrease with = lower doses=20 of Rsv and Quer. Surprisingly, 10 =B5M Rsv increased the MI = in the=20 first 24 h of treatment, but had no effect at later = time-points. Rsv=20 and Quer cotreatment induced a small reduction in MI, which was = not=20 observed by these treatments alone (Fig. 3c).=20 Cell cycle distribution indicated that Rsv (50 =B5M) caused a = modest G1=20 arrest (60 =B1 3% compared to 48 =B1 5% = control cells)=20 while Quer (100 =B5M) induced an S-phase accumulation=20 (20 =B1 3% compared to 16 =B1 2 control cells) = at=20 24 h. These effects were no longer observed at 72 h of = treatment=20 (data not shown).

=20

Reduction in pAkt by Rsv = and Quer is=20 not essential for their cytostatic effects. Given the = evidence=20 that PI3K/Akt is overactivated in a wide range of tumor types and = triggers=20 a cascade of responses that include cell growth, proliferation, = and=20 survival,⁽⁴⁵^{)=20
we wondered whether this pathway could be involved in Rsv and Quer =

apoptosis induction and growth arrest. Rsv (10 =B5M) and Quer =

(25 =B5M) reduced the phosphorylation of Akt to a level =
comparable to=20
the PI3K inhibitor LY without changing the intracellular Akt =
levels (Fig. 4a).=20
This effect was more intense within 30 min of treatment, but =
was also=20
observed at 4 h and even at 24 h (Fig. 4a).=20
This reduction in the phosphorylation of Akt was not due to =
alteration in=20
expression of PTEN, which was not affected by the treatments (Fig. 4b).}

=20

= Fig. 4. =20 Resveratrol (Rsv) and quercetin (Quer) decreased the=20 phosphorylation of Akt. C6 cells were treated with = 10 =B5M Rsv,=20 25 =B5M Quer, or 10 =B5M Rsv plus 25 =B5M = Quer, for=20 30 min, 4 h, and 24 h, and with 10 =B5M = LY294002=20 for 30 min. Lysates were subjected to western blot = analysis for=20 pS473Akt and total Akt antibodies (a) and PTEN antibody (b). = Representative blots of three independent experiments are = shown and=20 the numbers below the blots depict values of densitometric=20 evaluation as mean =B1 SEM. Number of C6 cells = treated as=20 indicated for 72 h (c). The results represent=20 mean =B1 SEM from three independent experiments. = anova was followed by SNK post = test.=20 *P < 0.05, **P < 0.01. =

[Normal=20 View ]

To verify if the reduction in the phosphorylation = of Akt was=20 responsible for the cell number decrease, we counted cells after = LY=20 treatment. While Rsv plus Quer reduced pAkt to 34% of the control = and the=20 cell number to 20% of the control, LY reduced the same parameter = to 26%=20 and 70% respectively (Fig. 4c).=20 Taken together, these results indicated that there was not a = relationship=20 between pAkt reduction and cell number decrease since LY was more = potent=20 in reducing pAkt and less effective in decreasing cell = number.

=20

Rsv and Quer cooperated in = inducing=20 senescence-like growth arrest in C6 cells. In order to = assess=20 a potential pro-senescence activity of chronically administered = Rsv and=20 Quer, we cultivated C6 cells in the presence of Rsv and Quer and=20 determined population doublings during 12 days. Rsv (10 =B5M) = or Quer=20 (25 =B5M) had only a small effect on the proliferation rate = of C6=20 cells, but Rsv plus Quer lead to a striking reduction in long-term = glioma=20 growth (Fig. 5a).=20 Senescence induction by Rsv plus Quer, measured by the activity of = acidic=20 β-galactosidase, a biochemical marker for lysosome=20 acidification,⁽⁴⁰^{)=20
was cooperative, since each treatment alone produced around 20% of =

SA-β-gal positive cells, whereas combination of Rsv and Quer =
induced=20
staining in almost all cells (Fig. 5b,c).}

=20

= Fig. 5. =20 Resveratrol (Rsv) and quercetin (Quer) induce = senescence in=20 C6 cells. C6 cells were cultivated in the continuous = presence of 10=20 or 25 =B5M Rsv, 25 =B5M Quer, 10 =B5M Rsv = plus 25 =B5M Quer,=20 or 10 =B5M LY294002 for 12 days and cumulative population = doublings=20 were plotted against time (a). After this time cells were = stained=20 for senescence-associated-β-galactosidade = (SA-β-gal) and=20 photographed (b). The bar graph represents the ratio of=20 SA-β-gal-positive cells to total cells of at least = three fields of=20 three independent experiments (c). Colony formation = efficiency of C6=20 cells 7 days after 12 days Rsv or Quer treatment or 6 days = of=20 cotreatment (d), and SA-β-gal staining at the end of = the colony=20 formation assay (e). anova was=20 followed by SNK post test. *P < 0.05, **P < 0.01, ***P < 0.001. =

[Normal=20 View ]

Despite a lack of relationship between pAkt = reduction and=20 cell number decrease in the acute treatments, we analyzed whether = Akt=20 played a role in chronic treatments. For this reason, we treated = cells for=20 12 days with 10 =B5M LY and determined population doublings. LY, = despite=20 blocking Akt (Fig. 4a),=20 did not affect the proliferation rate of C6 cells and these cells = did not=20 present SA-β-gal staining (Fig. 5a=96c),=20 corroborating the results found by acute treatments.

In order to assess the recovery ability of cells = that=20 survived the long-term Rsv and Quer treatments, we performed = colony=20 formation assay. For this, after 12 days of treatments we allowed = cells to=20 grow for 7 days without treatment and then we analyzed their = ability to=20 form colonies. The colony formation was reduced after treatment = with=20 25 =B5M Rsv, suggesting that in these cases, a sub-population = of cells=20 that survived the 12-day treatment were committed to arrest or = death, but=20 another sub-population survived and formed SA-β-gal-negative = colonies. (Fig. 5d,e).=20 At the end of 12 days of Rsv and Quer cotreatments, no cells = survived to=20 form colonies. Therefore, we performed a test after 6 days of = cotreatment=20 followed by 7 days without treatment and observed that the colony=20 formation was reduced by 40%; and as observed with the other = treatments,=20 cells that survived were not SA-β-gal-positive (Fig. 5d,e).

=20

=20

Discussion

In the present work we found that Rsv and Quer = elicited=20 cytostatic and cytotoxic effects on C6 glioma cell lines by = reducing cell=20 growth when administered acutely and chronically. The type of = effect=20 elicited was dependent on drug concentration and exposure time. = The=20 concentrations at/or above 50 =B5M Rsv induced necrosis, = apoptosis, and cell=20 cycle arrest in early time-points, whereas lower concentrations of = Rsv=20 required the presence of Quer for the induction of long-term = growth arrest=20 and senescence. Others have observed additive effects with acute=20 treatments with Rsv plus Quer in leukemia, breast cancer, and=20 melanomas,⁽^46=9648⁾=20 but these studies did not evaluate long-term effects of this=20 combination.

Inhibition of apoptosis and cell cycle = deregulation are two=20 mechanisms of tumor formation, and many chemopreventive agents act = through=20 the induction of apoptosis and cell cycle arrest to block the = carcinogenic=20 process.(^49=9652⁾=20 Our data showed that C6 cells treated with 50 =B5M Rsv for = 72 h=20 and Rsv plus Quer induced an increase in caspase 3/7 activity and = an=20 augmented annexin V staining. As soon as 24 h after = 50 =B5M Rsv=20 treatment, a strong decrease in MI accompanied by a small = reduction in the=20 G1 population was observed. These results support our notion that = the=20 cancer preventive effect of Rsv plus Quer may occur through the = induction=20 of apoptosis.(^8,53^{)=20
Previous works showed that the upregulation of p53-responsive =
genes such=20
as p21^WAF1/^CIP1, =
p300/CBP,=20
and Apaf1 by Rsv predisposed human prostate cancer (LNCaP) cells =
to=20
undergo apoptosis.⁽⁵⁴^{)=20
Similarly, Rsv- and Quer-induced apoptosis in HepG2 cells was =
accompanied=20
by a p53-dependent increase in Bax and =
p21.⁽^55,56^)<=
/SUP>}}

Cell cycle deregulation and apoptosis are closely = related=20 events. The PI3K/Akt pathway has a key role in regulating both = processes=20 and, as recently described, is involved in senescence=20 induction.⁽^57,58^{)<=
/SUP>=20
Besides this, it has been reported that increased PI3K activity =
due to=20
loss of PTEN, gene amplification, and mutation of catalytic and =
regulatory=20
subunits of PI3K is common in high grade=20
glioma,⁽⁵⁹^{)=20
and that inhibition of PI3K is cytotoxic in several human glioma =
cell=20
lines in vitro.⁽⁶⁰^{)=20
Both Rsv and Quer reduced the phosphorylation of Akt at early =
time-points=20
and despite reducing pAkt to a similar level to Rsv or Quer, LY =
produced=20
only a small reduction in cell numbers with acute and chronic =
treatments.=20
These results indicate that inhibition of the PI3K/Akt pathway =
does not=20
play an essential role in Rsv- or Quer-induced cell death. Other =
studies=20
have found an inhibitory effect of Rsv on the PI3K/Akt pathway, =
but in=20
this case the effect was mimicked by a PI3K=20
inhibitor.⁽⁶¹⁾}}}

The induction of senescence, irreversible growth = arrest, is=20 regarded in cancer cells as a means to halt tumor initiation and=20 progression.⁽⁶²^{)=20
Recent studies have revealed some molecular basis for cellular =
senescence,=20
which may involve the p38 MAPK cascade, and Rb and p53 tumor =
suppression=20
pathways.⁽⁶³^{)=20
Rsv and Quer are known for their antisenescent effects induced by =
a=20
variety of stresses, with only one report that shows Rsv-induced=20
senescence in the HCT116 colon cancer cell =
line⁽³¹^{)=20
and no report of Quer-induced senescence. The striking synergy =
observed=20
with low concentrations of Rsv and Quer point to the regulation of =

converging pathways to the senescence machinery.}}}

Long-term exposure to low concentrations of Rsv=20 (10=9625 =B5M) or Quer (25 =B5M) induced senescence-like = growth=20 arrest. More important, the clear additive =96 even synergistic = =96 effect=20 between Rsv and Quer was observed on cell number decrease, caspase = 3/7=20 activation, long-term proliferation inhibition, and senescence = induction.=20 This indicates that different targets may be involved in the = regulation of=20 these features and supports the notion that the combination of Rsv = and=20 Quer is an interesting strategy for glioma treatment.

Inhibition of Akt⁽⁵⁷^{)=20
as well as activation of Akt⁽⁵⁸^{)=20
have been linked to increase in senescence. Since we found that =
Rsv and=20
Quer reduced pAkt, we investigated whether these effects could be =
involved=20
with long-term growth arrest and senescence induction. LY, despite =

dramatically reducing Akt phosphorylation, did not have any effect =
on cell=20
growth and senescence, indicating that pAkt reduction is not =
solely=20
responsible for this process.}}

In this work we observed two types of effects, = with high=20 doses of Rsv inducing apoptosis, low doses inducing senescence, = and=20 cotreatment with low doses of Rsv and Quer inducing both. It is = still not=20 clear what determines if a cell will undergo senescence or = apoptosis.=20 Although most cells are capable of both, these processes seem to = be=20 mutually exclusive.⁽²⁸^{)=20
Cell type, gene expression, and level of tumour suppressor protein =

p53⁽^64,65^{)<=
/SUP>=20
appear to be determinant for the cell to undergo apoptosis or =
senescence.=20
Some studies clearly point to a crosstalk between the processes of =

apoptosis and cellular senescence;⁽⁶⁶^{)=20
however, the cross-regulation between these processes is far from=20
understood and further studies are needed to clarify their=20
relationship.}}}

The synergistic induction of senescence and = long-term growth=20 inhibition by Rsv and Quer suggests that the combination of these = agents=20 is a good candidate treatment for preclinical studies on glioma = tumors.=20 Studies in rodents indicate that both Rsv and Quer cross the = intact=20 blood=96brain barrier and reach concentrations in the range of 0.1 = and 0.3=20 =B5mol/kg, respectively.⁽^24,25^{)<=
/SUP>=20
Glioma growth may break the blood=96brain barrier, probably =
increasing the=20
concentration reachable inside the glioma. It is worthwhile noting =
that=20
Rsv is not toxic to primary astrocyte cultures and both Rsv and =
Quer are=20
protective against oxygen and glucose deprivation in hippocampal=20
organotypic cultures,⁽^7,17^{)=20
making these agents ideal for therapeutic intervention in CNS =
tumors.}}

Taken together, our studies add a novel facet to = the cancer=20 chemopreventive action of Rsv and Quer. Acutely administered high = doses of=20 Rsv can induce apotosis and chronically administered Rsv and Quer = in=20 subapoptotic doses can induce senescence-like growth arrest.

Several reports have investigated the effect of = individual=20 polyphenols. Because the majority of dietary compounds contain a = mixture=20 of polyphenols, it is important to understand their combinatorial = effect=20 at dietary concentrations. For a further potential application of = Rsv and=20 Quer, isolated or combined, as cancer chemopreventive agents or = adjuvants=20 in chemotherapy, more studies focusing on bioavailability, the = half-life=20 of Rsv and Quer, the interaction of both compounds and their = metabolites=20 are needed. This is even more necessary when taken into account = that=20 senescence induction normally requires the continuous presence of = the=20 inducing agents at the tumor site.

=20

=20

=20

Acknowledgments

This work was supported by the Brazilian funding = agencies=20 CNPq, FAPERGS, and PROPESQ/UFRGS. We thank Nicolle Barbieri for = help with=20 caspase assay and P=EDtia Ledur for help with preparation of the=20 manuscript.

=20

=20

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3D""=20

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