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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_01C96F28.6CA77AA0 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_01C96F28.6CA77AA0 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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"; =0A= } =0A= } =0A= }=0A= // set last two menu items to have no rollover and = a border at the top =0A= }=0A= menu.childNodes[x].childNodes[y].innerHTML =3D = tmpinnerhtml;=0A= // set default menu elements with diff style=0A= if (flag){=0A= if (more){=0A= liels[liels.length-3].style.borderTop =3D "1px solid = #DDDDDD";=0A= liels[liels.length-3].className =3D "noroll";=0A= liels[liels.length-2].style.borderTop =3D "1px solid = #DDDDDD";=0A= liels[liels.length-2].className =3D "noroll";=0A= liels[liels.length-1].className =3D "noroll";=0A= more =3D false;=0A= } else {=0A= liels[liels.length-2].style.borderTop =3D "1px solid = #DDDDDD";=0A= liels[liels.length-2].className =3D "noroll";=0A= liels[liels.length-1].className =3D "noroll";=0A= } =0A= flag =3D false; =0A= } =0A= =0A= // set sub menu title for the msnu=0A= liels[0].className =3D "hoverTitle";=0A= // apply rollover functions to hide/show sub menus=0A= if (classname =3D=3D "menuDOM"){=0A= menu.childNodes[x].onmouseover =3D function(){=0A= // safari doesnt handle auto in absoulte = positioning correctly so this code makes the menus display correctly.=0A= if (navigator.userAgent.indexOf("Safari") !=3D -1){=0A= if(!window.devicePixelRatio) {=0A= this.getElementsByTagName("UL")[0].style.marginTop =3D "20px";=0A= this.getElementsByTagName("UL")[0].style.marginLeft =3D "0";=0A= }=0A= }=0A= this.className =3D"menuhover domArrowOver";=0A= if(document.all) = this.getElementsByTagName("iframe")[0].style.height=3Dthis.getElementsByT= agName("ul")[0].offsetHeight;=0A= }=0A= menu.childNodes[x].onmouseout=3Dfunction() {=0A= this.className =3D "domArrowOut";=0A= }=0A= }=0A= =0A= if(document.all) {=0A= var newIframe =3D document.createElement("iframe");=0A= newIframe.style.filter =3D = "progid:DXImageTransform.Microsoft.Alpha(style=3D0,opacity=3D0)";=0A= menu.childNodes[x].appendChild(newIframe);=0A= }=0A= =0A= } =0A= }=0A= // deletes the sub menus on the IEMAC as they have a = background bug, will still remain on NS6=0A= if ((navigator.userAgent.indexOf("Mac") !=3D -1) && = (navigator.userAgent.indexOf("MSIE") !=3D -1)){=0A= menu.childNodes[x].innerHTML =3D = menu.childNodes[x].innerHTML.split(""=0A= var menu =3D document.all["menu"].children[0];=0A= menu.style.marginBottom =3D "-3px";=0A= for (var x=3D1; 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Syst= em=20 Maintenance: Jan 2 2009

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Brain=20 Pathology

ISSN: 1750-3639 (Online)

ISSN: 1015-6305 (Print)

Volume=20 19, Issue 1, 2009.

Pages: 121=96131

Tables: 1,=20 2,=20 3

List of=20 Images

Abstract=20 Only

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3D"About

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

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

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

'); } =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('

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');}}=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('Figure 1. Astrocytomas in = NPcis mice. A. Shows diffuse dysplastic nuclei in a World = Health Organization grade (WHO) II astrocytoma. Approximately 40% of = astrocytomas observed in NPcis mice are WHO II, depending of the = genetic background. B. Shows an anaplastic astrocytoma in the = spinal cord (V). Approximately 50% of astrocytomas observed are WHO III = in the brain or spinal cord. Up to 40% of astrocytomas are found in the = spinal cord and many are suggestive of a primary spinal cord lesion, as = opposed to infiltration from a primary brain tumor. C. Shows an = aggressive glioblastoma multiforme (GBM) that appears to have exited the = brain at the bottom of the panel, broken through the skull (S), and is = spreading along the surface of the skull at the top of the panel. = D.,E. Show examples of diagnostic criteria in NPcis = astrocytomas. Dysplastic nuclei are seen in all astrocytomas (D) = with distinctive multinucleated giant cells (arrows) found in up to 15% = of astrocytomas, including most WHO IV tumors. WHO III and WHO IV = astrocytomas have varying degrees of mitotic activity (E). Arrows = point at a couple of the mitoses visible in the panel. (F) WHO IV = GBMs have regions of N. Scale bars indicate 100 µm. = V =3D vertebra; S =3D skull; = N =3D necrosis.'); }=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('Figure 2. Secondary structures = found in NPcis astrocytomas. A. Shows an example of = satellitosis in which tumor cells form satellite structures around large = neurons. An especially distinctive satellitosis pattern is found in up = to 7% of observed astrocytomas. B. Shows an example of = rosette-like structures (*) in which tumor cells cluster and fan out = from a central point. C. Shows an example of perivascular = structures in which tumor cells line up along blood vessels in close = association. D. Shows an example of hemorrhage (H) that is found = in most GBMs and many of the more aggressive anaplastic astrocytomas. = E.,F. Show examples of pseudopallisading tumor cells = around a central necrotic core (N) that is found in rare cases of = GBM. Scale bars indicate 100 µm. = H =3D hemorrhage.'); }=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('Figure 3. Genetically engineered = murine peripheral nerve sheath tumors in NPcis mice. = A.,B. SC, DRG and N are indicated for different MPNSTs. = D. Shows a high magnification field of the spinal nerve root = tumor shown in C. (box), and the arrow points to a mitotic figure = found in the tumor. Scale bars indicate 100 µm. = SC =3D spinal cord; DRG =3D dorsal root ganglion; = N =3D nerve.'); }=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('Figure 4. Identifying genetic = risk factors between susceptible and resistant strains using F2 = intercross or backcross designs. To identify a dominant modifier of = resistance (M), a susceptible strain and a resistant strain are crossed = to generate F1 progeny. Germline recombination events in F1 progeny will = generate chromosomes that are a mixture of the two strain backgrounds. = F1 progeny can be intercrossed to give rise to F2 progeny carrying = different recombination events on each chromosome and allowing the = possibility of detective recessive modifiers. Alternatively, F1 progeny = can be backcrossed to the strain carrying the recessive allele of the = modifier, simplifying the detection of dominant modifiers.'); }=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('Figure 5. Reference panels for = identifying genetic risk factors. Panels of strains with stable genetics = and well-characterized genotypes can be used to identify modifier genes. = CSS carry one chromosome from one strain (black) on the background of = another strain (gold). By examining the phenotype in a CSS line for each = autosomal chromosome, the X and Y chromosomes, and the mitochondria, one = can isolate which chromosomes give rise to genetic variation in = phenotype. Recombinant inbred strains carry different mixtures of two = strains (black and gold), with different combinations in each line. By = examining a phenotype across a large number of these lines, one can = identify which regions of the genome associate with a particular trait. = Because there is limited variation between any two inbred strains, = higher order heterogeneous stocks have been generated. A recent example = of this is the CC in which eight different strains are combined to = preserve variability and then inbred to form lines, with each line = carrying different combinations of the eight parental lines. = CSS =3D chromosome substitution strains; CC =3D = collaborative cross; RXS =3D resistant X susceptible = recombinant inbred line.'); }=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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'); doctail(hwnd); = hwnd.document.close(); hwnd.focus(); } ------=_NextPart_000_0000_01C96F28.6CA77AA0 Content-Type: application/octet-stream Content-Transfer-Encoding: quoted-printable Content-Location: http://www3.interscience.wiley.com/cgi-bin/fulltext/121543785/main.html,ftx_abs Wiley InterScience = :: Article Full Text HTML setDOI("ADOI=3D10.1111/j.1750-3639.2008.00236.x");<= /SCRIPT>

3D""=20

Brain = Pathology

=20 What=20 is RSS?

Volum= e=20 19, Issue 1, Pages 121-131

Published Online: 3 Dec 2008

3D"E-Mail

3D""=20

< Previous=20 Article | Next=20 Article = >

Save=20 Article to My Profile=20 Download=20 Citation=20

Abstract=20 | References=20 | Full Text: HTML , PDF=20 (1430k)=20

View=20 Full Width

3D""=20

3D""=20

MINI-SYMPOSIUM: Mouse Models of Brain=20 Tumors

Brain Tumor Susceptibility: the Role = of Genetic=20 Factors and Uses of Mouse Models to Unravel Risk

Karlyne=20 M. Reilly, Ph.D.

Mouse Cancer Genetics = Program,=20 National Cancer Institute-Frederick, Frederick, Md.

Correspondence to Karlyne = M. Reilly,=20 PhD, Mouse Cancer Genetics Program, National Cancer = Institute-Frederick,=20 West 7th Street at Fort Detrick, PO Box B, Frederick, MD 21702 = (E-mail: kreilly@ncifcrf.gov)=20

Copyright Journal compilation =A9 2009 = International Society of Neuropathology

KEYWORDS

astrocytoma =95 genetic risk factors =95 = glioblastoma=20 multiforme =95 mouse models =95 neurofibromatosis =95 peripheral = nerve sheath=20 tumors

ABSTRACT

3DAbstract=20

=

=

=

=

<= /TD>

=

Brain tumors are relatively rare but deadly = cancers, and=20 present challenges in the determination of risk factors in the = population.=20 These tumors are inherently difficult to cure because of their = protected=20 location in the brain, with surgery, radiation and chemotherapy = options=20 carrying potentially lasting morbidity for patients and incomplete = cure of=20 the tumor. The development of methods to prevent or detect brain = tumors at=20 an early stage is extremely important to reduce damage to the = brain from=20 the tumor and the therapy. Developing effective prevention or = early=20 detection methods requires a deep understanding of the risk = factors for=20 brain tumors. This review explores the difficulties in assessing = risk=20 factors in rare diseases such as brain tumors, and discusses how = mouse=20 models of cancer can aid in a better understanding of genetic risk = factors=20 for brain tumors.

Received 22 September 2008; accepted 7 October=20 2008.

DIGITAL OBJECT IDENTIFIER (DOI)

10.1111/j.1750-3639.2008.00236.x About = DOI

=20

=20

=20

ABSENCE OF EVIDENCE IS NOT EVIDENCE = OF=20 ABSENCE

=

=

=

=

<= /TD>

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The debate over the past decade on whether cell = phone use=20 increases brain tumor risk highlights the difficulties in = identifying risk=20 factors for brain tumorigenesis. Multiple studies have produced=20 conflicting results, and although it is now accepted that the risk = of=20 developing a brain tumor from cell phone use is likely negligible = (1,=20 25, 34, 51, 66), the few studies showing an effect of cell = phones=20 continue to raise concerns in the general population. Although = risk=20 factors and causes of many common cancers have now been = established (ht= tp://www.cancer.gov/cancertopics/prevention-genetics-causes),=20 understanding the factors that contribute to brain tumors remains=20 elusive.

=20

=20

DISEASE RISK FACTORS

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There are multiple types of risk factors governing = disease=20 susceptibility. These include behavioral risk factors, such as = choosing to=20 smoke or excessive alcohol consumption, environmental risk = factors, such=20 as exposure to chemical carcinogens or radiation, biological risk = factors,=20 such as puberty or aging and genetic risk factors, such as = inheritance of=20 tumor suppressor mutations or susceptibility alleles of modifier = genes.=20 These different risk factors form a spectrum of what people can = control to=20 what they cannot control, with behavioral risk factors being the = easiest=20 to control and biological risk factors being impossible to = control.=20 Furthermore, different types of risk factors interact. For = example, the=20 behavioral risk factor of smoking likely interacts with genetic=20 variability in the population that determines how easily a person = becomes=20 addicted to cigarettes. Similarly, wearing sunscreen is a = behavioral=20 factor that reduces risk, counteracting the environmental risk of = UV=20 exposure. By understanding different types of risk factors, it is = hoped=20 that protective factors that are controllable (such as wearing = sunscreen)=20 can be developed to counteract risk factors that are = uncontrollable (such=20 as UV exposure). In the case of brain tumors, this might include = increased=20 screening for early detection of tumors in individuals at higher = risk=20 because of genetic susceptibility. While an individual's exposure = to=20 behavioral and environmental risk can be difficult to study and = can change=20 at different times of life, genetic risk factors are expected to = be stable=20 in the individual over their lifetime. This article will focus = primarily=20 on genetic determinants of brain tumor risk because an = individual's=20 genetic background forms the foundation against which all other = risk=20 factors interact.

=20

=20

GENETIC RISK FACTORS FOR BRAIN = CANCER

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Although very little is known about the genetic = risk factors=20 for brain cancer, a few factors have been identified thus far. = Brain=20 tumors are associated with several familial cancer predisposition=20 syndromes. These include Li-Fraumeni syndrome, neurofibromatosis, = tuberous=20 sclerosis and Turcot's syndrome. In these syndromes, individuals = inherit a=20 germline mutation in a tumor suppressor gene. Tumors initiate when = the=20 remaining copy of the tumor suppressor is mutated or silenced, = giving rise=20 to cells with a growth advantage. Because tumorigenesis requires = the=20 accumulation of multiple mutations in cells, these individuals = have an=20 increased tumor risk because all cells carry an initial mutation.=20 Li-Fraumeni syndrome is caused by mutations in the cell checkpoint = genes=20 TP53(40)=20 and CHEK2(4).=20 Turcot's syndrome is caused by mutations in genes involved in DNA = repair=20 (24).=20 It is likely that in Li-Fraumeni syndrome and Turcot's syndrome = the risk=20 for brain tumors is increased by an increased rate of DNA mutation = leading=20 to uncontrolled growth. Neurofibromatosis is caused by mutations = in NF1 or NF2(20,=20 29) and tuberous sclerosis is caused by mutations in TSC1 or TSC2(2,=20 36). NF1, NF2, TSC1 and=20 TSC2 are all involved in down regulation of = growth=20 promoting signal transduction pathways in the cell. It is = therefore likely=20 that in neurofibromatosis and tuberous sclerosis, the risk for = brain=20 tumors is increased because brain cells are primed for excessive = growth=20 and then develop additional mutations allowing cancer to form.

In addition to known familial cancer = predisposition=20 syndromes, it has also been observed that brain tumors can cluster = within=20 families [see (45)=20 for review]. Familial clustering can be because of both genetic = and=20 environmental factors, as families often share common = environmental=20 exposures in addition to common genes. Modeling of the inheritance = pattern=20 of familial glioma suggests that at least in some cohorts, genetic = factors=20 play a role in susceptibility (15,=20 41, 42). Segregation analysis of 297 families in Sweden = suggested that=20 recessive genes may contribute to familial glioma, although a=20 multifactorial model was not excluded (41).=20 However, homozygosity mapping in a study of seven glioma families = in=20 Sweden did not identify any common homozygous alleles (44).=20 Segregation analysis of 639 families in the United States and = Canada found=20 evidence for a multifactorial Mendelian model, and suggested that = familial=20 glioma is affected by multiple low penetrance genes (15).=20 In contrast, a study of 396 families in Iceland found no evidence = for=20 increased risk of gliomas in families, although the authors cite = several=20 limitations to the study including small sample size (53).

Although evidence for a genetic risk component in = brain=20 tumors has been demonstrated, efforts to identify the underlying = genes=20 have had little success. A study of seven families in Sweden found = weak=20 linkage to chromosome 1q23 (logarithm of the odds=20 (LOD) =3D 1.0517) (44).=20 In a study of 15 families in Finland, linkage for susceptibility = to glioma=20 was found on four chromosomes, 1q25.1, 6q27, 8p21.3 and 15q26.2, = with=20 chromosome 15 showing the most significant linkage. Because the = linked=20 region on chromosome 15 covers 40 cM, it was not possible to = identify=20 genes in this study (58).=20 These studies are severely hampered by the fact that brain tumors = are rare=20 and different brain tumor types often need to be pooled together = in=20 studies, possibly diluting out significant effects on specific = brain tumor=20 histologies. To begin to address these issues, the GLIOGENE = international=20 consortium has recently formed to collect large numbers of brain = tumor=20 cases (45).

Because of the difficulties in using unbiased = screens for=20 genetic risk factors, many investigators have turned to candidate=20 approaches to test for susceptibility genes. One of the earliest = gene=20 families to be examined was the glutathione S-transferase (GST) = family of=20 genes because of their role in detoxifying environmental = carcinogens.=20 Studies attempting to link specific GST polymorphisms to brain = tumor risk=20 have given conflicting results, with most studies finding no = significant=20 link between GST genotype and brain tumor risk (37,=20 54, 69, 73). Patients carrying a combination of the germ-line = genotype=20 GSTP1*A/*A=20 and GSTM1 null were shown to have an = increased=20 survival, but also have greater adverse effects in response to=20 chemotherapy (54).=20 A second candidate approach has been to examine the role of genes = involved=20 in DNA repair, such as p53 and XRCC (6,=20 35, 43, 46, 77). These studies have also shown mixed results, = with=20 polymorphisms in p53 showing increased risk in glioblastoma in one = study=20 (46),=20 but not another (77),=20 and polymorphisms in XRCC7 being associated = with risk=20 for glioma (77).=20 Recently, a polymorphism in the apoptosis gene Caspase=20 8 has also been shown to be associated with increased = glioma risk,=20 although in four out of the five case control studies examined, = this=20 association was not statistically significant (5).=20 An observed inverse relationship between asthma and glioma (81)=20 has led investigators to examine polymorphisms in genes associated = with=20 immune response and inflammation. Studies of the association of=20 polymorphisms in IL13 and IL4R with glioblastoma risk have = produced=20 conflicting results (67,=20 68, 80).

The studies to link candidate genes to brain tumor = risk=20 overall have yielded mixed results, pointing out the difficulty of = understanding the genetic basis of brain tumor risk, even when = evidence=20 exists for this genetic risk being important. The difficulty in = finding=20 genetic risk factors stems from the limited number of brain tumor = patients=20 available for these studies, the possibility for necessary = interactions of=20 different genetic factors, such that a single candidate factor = does not=20 show significant linkage, and the possibility that genetic factors = interact with environmental risk factors that are not uniform = across=20 different populations and different studies. In essence, this = boils down=20 to too many variables and not enough subjects to determine = statistical=20 significance. While the GLIOGENE consortium seeks to build a large = study=20 population, this may not entirely solve the problem if confounding = variables and heterogeneity in the population, both at the genetic = and=20 environmental level, are the issue.

=20

=20

MOUSE MODELS OF BRAIN CANCER TO = DISSECT ISSUES=20 OF HETEROGENEITY

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When studying genetic risk factors in human = populations, one=20 must consider and control for many confounding variables. Mouse = models of=20 human cancer allow one to start with a relatively homogeneous = system and=20 layer on complexity in a controlled way. This allows one to = determine=20 genetic risk factors in an unbiased screen using fewer subjects = than would=20 be possible in a human study. Genetic risk factors that are = identified in=20 mice can then be tested for association in human populations using = more=20 straightforward candidate approaches. Mouse models of brain cancer = can be=20 bred under relatively constant environmental conditions with a = constant=20 diet and a well-controlled genetic background to isolate genetic = factors=20 from environmental factors. Once the genetic factors are well = understood,=20 environmental factors can be introduced into the experiment, for = example=20 by feeding mice a high-fat diet or exposing them to pathogens or=20 carcinogens, to determine the effects on cancer risk. Mouse models = of skin=20 cancer, lung cancer, colon cancer, plasmocytoma, teratoma, and = breast=20 cancer metastasis have already been used to identify modifier = genes of=20 cancer susceptibility (13,=20 16, 19, 21, 55, 64, 84, 85). Ongoing experiments are using a = mouse=20 model of astrocytoma and secondary glioblastoma to identify = modifiers of=20 brain cancer (61).

The power of mapping modifiers of brain tumors in = mouse lies=20 in the ability to generate large numbers of homogeneous = individuals. A=20 mouse model of astrocytoma and secondary glioblastoma has been = generated=20 by combining mutations in Nf1 and Trp53 on the same chromosome of the mouse (60).=20 Because the mutations are tightly linked on the same chromosome, = they are=20 inherited as a single mutation, resulting in near Mendelian = inheritance=20 ratios. These mice develop brain tumors with high penetrance in = the=20 population, such that in a population of mutant progeny, up to 70% = of=20 animals will develop a brain tumor (Figure 1)=20 with many of the characteristics of human astrocytoma and = glioblastoma (Figures 1=20 and 2). This combination of high penetrance and simple = genetics means=20 that large numbers of mice developing brain tumors can be = generated=20 relatively easily for studies of brain tumor risk.

=20

= Figure 1. Astrocytomas in NPcis mice.=20 A. Shows diffuse dysplastic nuclei in = a World=20 Health Organization grade (WHO) II astrocytoma. = Approximately 40% of=20 astrocytomas observed in NPcis mice = are WHO II,=20 depending of the genetic background. B. Shows=20 an anaplastic astrocytoma in the spinal cord (V). = Approximately 50%=20 of astrocytomas observed are WHO III in the brain or spinal = cord. Up=20 to 40% of astrocytomas are found in the spinal cord and many = are=20 suggestive of a primary spinal cord lesion, as opposed to=20 infiltration from a primary brain tumor. C.=20 Shows an aggressive glioblastoma multiforme (GBM) that = appears to=20 have exited the brain at the bottom of the panel, broken = through the=20 skull (S), and is spreading along the surface of the skull = at the=20 top of the panel. D.,E.=20 Show examples of diagnostic criteria in NPcis=20 astrocytomas. Dysplastic nuclei are seen in all astrocytomas = (D) with distinctive multinucleated giant = cells=20 (arrows) found in up to 15% of astrocytomas, including most = WHO IV=20 tumors. WHO III and WHO IV astrocytomas have varying degrees = of=20 mitotic activity (E). Arrows point at = a couple=20 of the mitoses visible in the panel. (F) WHO IV=20 GBMs have regions of N. Scale bars indicate 100 =B5m.=20 V =3D vertebra; S =3D skull;=20 N =3D necrosis.

[Normal View=20 ]

=20

= Figure 2. Secondary structures found in NPcis astrocytomas. A. = Shows an=20 example of satellitosis in which tumor cells form satellite=20 structures around large neurons. An especially distinctive=20 satellitosis pattern is found in up to 7% of observed = astrocytomas.=20 B. Shows an example of rosette-like = structures=20 (*) in which tumor cells cluster and fan out from a central = point.=20 C. Shows an example of perivascular = structures=20 in which tumor cells line up along blood vessels in close=20 association. D. Shows an example of = hemorrhage=20 (H) that is found in most GBMs and many of the more = aggressive=20 anaplastic astrocytomas. E.,F. Show examples of pseudopallisading tumor = cells=20 around a central necrotic core (N) = that is=20 found in rare cases of GBM. Scale bars indicate = 100 =B5m.=20 H =3D hemorrhage.

[Normal=20 View ]

The Nf1 gene encodes the = protein=20 neurofibromin that acts as a rasGAP protein to downregulate active = ras=20 signaling (3,=20 22, 26, 47, 82, 83). The gene is mutated in the disease NF1 in = which=20 patients develop many benign tumors of the nervous system and are = at=20 increased risk for developing astrocytomas, glioblastomas and = malignant=20 peripheral nerve sheath tumors. Nf1 has = recently been=20 shown to be mutated in a subset of human glioblastoma samples (48,=20 57). Trp53 encodes the p53 protein that = acts as a=20 regulator of cell cycle arrest and apoptosis. The human gene for = p53=20 (Tp53) is mutated in the disease = Li-Fraumeni syndrome=20 in which patients are at an increased risk for developing many = types of=20 cancer, including brain tumors. Tp53 or = other members=20 of the p53-signaling pathway are frequently mutated in human = astrocytoma=20 and glioblastoma samples. Both Nf1 and = Trp53 have been mutated in mice. Homozygous = mutation of=20 Nf1 is embryonic, lethal and heterozygous = mutation of=20 Nf1 is tumor-prone (8,=20 32). Homozygous mutants for Trp53 = develop thymic=20 lymphoma with high penetrance, whereas heterozygous mutants = develop a=20 variety of tumor types, primarily sarcomas (18,=20 31). When Nf1 and Trp53=20 mutations are combined on the same chromosome through a germline=20 recombination event, the resulting Nf1−/+;Trp53−/+cis(NPcis) mice develop malignant peripheral nerve = sheath=20 tumors and astrocytomas with high penetrance (11,=20 75).

Studies of the NPcis mouse = model on=20 different strain backgrounds have demonstrated the importance of = genetic=20 risk factors in astrocytoma and glioblastoma, as well as the = complexity of=20 interactions between different types of risk factors. NPcis mice on a C57BL/6J (B6) strain background = are highly=20 susceptible to astrocytomas and glioblastomas, with up to 70% of = males=20 developing brain tumors (Table 1).=20 In contrast, NPcis mice on a 129S4/SvJae = (129S4)=20 strain background are highly resistant to developing astrocytoma = and=20 glioblastoma, with as few as 12% of females developing brain = tumors (Table 1).=20 In addition to the effect of strain background, the inheritance of = the=20 NPcis mutant chromosome from the mother or = father has=20 an effect on astrocytoma risk. In the case of females, progeny of = NPcis mutant mothers develop higher-grade tumors = than=20 progeny of mutant fathers, although the percentage of the = population with=20 brain tumors is the same. In the case of males, progeny of NPcis mutant mothers develop more tumors than = progeny of=20 mutant fathers (Table 1).=20 This data demonstrates that there is a parent-of-origin effect on = brain=20 tumors, likely because of changes in expression of an imprinted = gene on=20 mouse chromosome 11. Interestingly, this imprinted effect has=20 gender-specific consequences, with females developing higher-grade = tumors=20 and males developing more brain tumors in the population.

=20

Table 1. Variation in=20 astrocytoma incidence in NPcis mice depends = on strain=20 background.

Cross	Females (%)	Males (%)

NPcis B6 mother=20 X WT B6 father	51	71
WT B6 mother X NPcis B6 father	52	49
WT 129S4 mother X NPcis 129S4 father	12	17

WT =3D wild-type.

Studies are ongoing to map the modifier loci = responsible for=20 the difference in susceptibility between the B6 and 129S4 strains. = However, comparisons between F1 intercrosses of NPcis=20 mice on a 129S4 background and NPcis mice = on the B6=20 background have identified a modifier locus on mouse chromosome 11 = in a=20 30-Mb congenic region surrounding the Trp53 = and Nf1 mutations (61).=20 It is interesting to note that there are strain-specific = differences in=20 the expression level of Nf1 in the brain on = the B6=20 and 129S4 strain backgrounds (27).=20 It is therefore possible that levels of Nf1 = expression in the heterozygous brain cell, before the wild-type = (WT) copy=20 of Nf1 is lost, could alter the risk of = developing=20 astrocytoma. Further studies are necessary to confirm or disprove = this=20 possibility. In addition, there are multiple candidate genes in = the region=20 that are polymorphic between B6 and 129S4, and could be = responsible for a=20 change in brain tumor risk (61).

In addition to using the NPcis mouse=20 model to identify genetic risk factors for brain tumors, the model = has=20 also been used to examine genetic risk factors for malignant = peripheral=20 nerve sheath tumors [genetically engineered murine peripheral = nerve sheath=20 tumors (GEM PNSTs), in mouse (71)]=20 (Figure 3)=20 (62).=20 These studies have been very informative on the complexity of = genetic=20 interactions contributing to nervous system tumor risk, and are = shaping=20 the design of experiments to understand risk of brain tumors. The = risk of=20 GEM PNSTs is also affected by inheritance of the NPcis mutant chromosome from the mother or the = father. In=20 contrast to what is seen with brain tumors, the progeny of mutant = fathers=20 have an increased incidence of GEM PNSTs compared to the progeny = of mutant=20 mothers (Table 2),=20 with males having a greater risk of GEM PNSTs than females = regardless of=20 the cross. Crosses of NPcis B6 mice to the = A/J strain=20 have demonstrated the existence of dominant resistance alleles in = A/J and=20 backcross mapping has been used to identify loci responsible for=20 resistance to GEM PNSTs. Two nerve sheath tumor resistance loci, = Nstr1 and Nstr2, have been = identified=20 thus far (62),=20 and the ability of these loci to affect tumor resistance is cross=20 specific. Nstr1 on mouse chromosome 19 only = appears=20 to affect tumor resistance in progeny of NPcis mutant=20 fathers, whereas Nstr2 on mouse chromosome = 15 appears=20 to affect tumor resistance only in progeny of NPcis=20 mutant mothers. When the combined population of progeny of mutant = fathers=20 and progeny of mutant mothers were examined for resistance loci, = no=20 significant loci were found. This is due to the fact that because = the=20 resistance loci only act in a subset of the population, the other=20 unaffected individuals mask the ability to identify significant = linkage.=20 The heterogeneity of the population, even within this well-defined = mouse=20 population, prevents the identification of genetic risk factors = that are=20 present in the population. This further illustrates the = difficulties for=20 identifying genetic risk factors directly in patient populations, = where=20 the degree of heterogeneity is far greater.

=20

= Figure 3. Genetically = engineered=20 murine peripheral nerve sheath tumors in NPcis=20 mice. A.,B. SC, DRG and N are indicated for = different MPNSTs.=20 D. Shows a high magnification field = of the=20 spinal nerve root tumor shown in C. = (box), and=20 the arrow points to a mitotic figure found in the tumor. = Scale bars=20 indicate 100 =B5m. SC =3D spinal cord;=20 DRG =3D dorsal root ganglion; = N =3D nerve.=20

[Normal View=20 ]

=20

Table 2. Variation in=20 genetically engineered murine peripheral nerve sheath tumor = incidence in=20 NPcis mice depends on parental = inheritance.=20

Cross	Females (%)	Males (%)

NPcis B6 mother=20 X WT B6 father	34	46
WT B6 mother X NPcis B6 father	60	82

WT =3D wild-type.

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IDENTIFYING MOUSE GENETIC RISK = FACTORS FOR=20 CANCER IN THE 21ST CENTURY

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Since the sequencing of the human (38,=20 65, 74) and mouse genomes (39,=20 50, 76, 78) at the beginning of this century, there has been a = rapid=20 acceleration in the ability to map genetic risk factors. These = efforts=20 have led to the identification of vast numbers of single = nucleotide=20 polymorphisms for following genetic variability in both human and = mouse,=20 and made possible the genome-wide association studies currently = underway=20 in patient populations (59).=20 In parallel, new research tools being developed in the mouse = genetics=20 community are expected to provide new power to identifying genetic = risk=20 factors in mice.

Thus far, most of the modifiers of cancer = identified in mice=20 have been found through a combination of mapping in backcross = populations,=20 intercross populations (Figure 4),=20 recombinant congenic strains as well as candidate gene approaches = (7,=20 12, 13, 19, 21, 55, 64, 84, 85). These approaches take = advantage of=20 the relative simplicity of comparing two strains through different = genomic=20 combinations to find regions of the genome responsible for = determining=20 cancer risk. These approaches face the challenges of needing to = genotype a=20 large number of mice at a large number of genetic loci in order to = correlate the change in phenotype to a particular genotype at a = particular=20 point in the genome. Although the ability to genotype large = numbers of=20 loci is becoming less costly, efforts have been made to develop = strain=20 tools that are fixed in their genotype and therefore do not = require=20 genotyping for each experiment performed. These include = recombinant inbred=20 strain panels and chromosome substitution strains (Figure 5)=20 (52,=20 70). These panels have the advantage that they are genetically = stable=20 and so can be compared across different experiments. As an = example, if one=20 found certain strain affected one kind of brain tumor, one could = easily go=20 back the these strains and test for effects on other types of = brain=20 tumors. More advanced strain tools are now being employed to = further=20 balance the need for a simple, experimentally tractable system = with the=20 need to increase the heterozygosity of the system to improve = mapping=20 resolution [see (28)=20 for a recent review].

=20

= Figure 4. Identifying = genetic risk=20 factors between susceptible and resistant strains using F2=20 intercross or backcross designs. To identify a dominant = modifier of=20 resistance (M), a susceptible strain and a resistant strain = are=20 crossed to generate F1 progeny. Germline recombination = events in F1=20 progeny will generate chromosomes that are a mixture of the = two=20 strain backgrounds. F1 progeny can be intercrossed to give = rise to=20 F2 progeny carrying different recombination events on each=20 chromosome and allowing the possibility of detective = recessive=20 modifiers. Alternatively, F1 progeny can be backcrossed to = the=20 strain carrying the recessive allele of the modifier, = simplifying=20 the detection of dominant modifiers.

[Normal=20 View ]

=20

= Figure 5. Reference panels = for=20 identifying genetic risk factors. Panels of strains with = stable=20 genetics and well-characterized genotypes can be used to = identify=20 modifier genes. CSS carry one chromosome from one strain = (black) on=20 the background of another strain (gold). By examining the = phenotype=20 in a CSS line for each autosomal chromosome, the X and Y=20 chromosomes, and the mitochondria, one can isolate which = chromosomes=20 give rise to genetic variation in phenotype. Recombinant = inbred=20 strains carry different mixtures of two strains (black and = gold),=20 with different combinations in each line. By examining a = phenotype=20 across a large number of these lines, one can identify which = regions=20 of the genome associate with a particular trait. Because = there is=20 limited variation between any two inbred strains, higher = order=20 heterogeneous stocks have been generated. A recent example = of this=20 is the CC in which eight different strains are combined to = preserve=20 variability and then inbred to form lines, with each line = carrying=20 different combinations of the eight parental lines.=20 CSS =3D chromosome substitution strains; = CC =3D =20 collaborative cross; RXS =3D resistant X = susceptible=20 recombinant inbred line.

[Normal=20 View ]

Because most mapping of genetic risk factors in = mouse=20 focuses on the comparison between two inbred strains, there are=20 limitations to the number of modifiers that can be identified. = Despite the=20 differences between the inbred strains, there is relatively low = allelic=20 diversity between strains (63).=20 This is because of the fact that the laboratory inbred strains = were=20 developed only recently, thus their evolutionary history is short. = To=20 address this issue, mouse geneticists are developing an expanded = panel of=20 recombinant inbred lines, termed the Collaborative Cross, which is = derived=20 from eight different founder strains (10,=20 72). The eight founder strains were chosen for genetic and = phenotypic=20 diversity between different strains and subspecies, allowing for = the=20 capture of close to 90% of the existing variation in the mouse = genome (63)=20 (Figure 5).=20 Through a collaboration of three different breeding locations, up = to 1000=20 independent Collaborative Cross lines are planned and currently = being=20 generated (9,=20 30, 49). These lines will be densely genotyped, such that by = comparing=20 phenotypes across many lines one can map modifiers precisely. = Modeling of=20 the distribution of allelic frequency in the Collaborative Cross = suggests=20 that it closely mimics the distribution found in the human = population (63).=20 This resource should allow a system genetics approach to = understanding=20 genetic risk to disease, and allow researchers to examine = interactions at=20 multiple loci in a controlled way.

Characterized strains of wild-type mice do not = develop=20 spontaneous brain tumors with high enough frequency or short = enough=20 latency to provide a tractable experimental system for mapping = genetic=20 risk factors. It is therefore important to choose appropriate = genetically=20 engineered mouse models for mapping studies. Mouse models on = well-defined=20 strain backgrounds can be crossed into backcross or intercross = mapping=20 designs (Figure 4)=20 by maintaining the engineered mutations at each generation, or can = be=20 crossed to genetic reference panels such as shown in Figure 5=20 to look at F1 hybrids for genetic risk factors that act dominantly = over=20 the strain background of the brain tumor model. These experiment = are best=20 performed with "one-allele/one-generation" models. These are = models in=20 which the brain tumor phenotype can be observed in the = heterozygous state,=20 and either require the inheritance of only one engineered allele, = or in=20 which multiple alleles can be homozygosed in a parental line and = crossed=20 to the reference panels to examine the phenotype in heterozygotes. = Mouse=20 model systems in which many engineered alleles need to be = combined, such=20 as in the case for many Cre-flox models, will be difficult to use = for=20 mapping studies because of the low rate of co-inheritance of = multiple=20 alleles in mouse crosses. Several mouse models are available that = fit the=20 one allele/one generation criteria, shown in Table 3,=20 although not all of these may be currently available on an inbred = strain=20 background. The NPcis model acts as a = one-allele=20 model, because although two genes are mutated in the model, they = are=20 tightly linked on the same chromosome and so are inherited as a = single=20 allele. Inbred mouse models can be directly crossed to genotyped = genetic=20 reference panels to determine the location of modifier loci = without=20 further genotyping, since all of the genotypes of the progeny can = be=20 inferred from the parents.

=20

Table 3. Examples of=20 "one allele/one generation" mouse models for mapping brain tumor = genetic=20 risk factors in genetic reference populations. =

Mouse model	Brain tumor	Penetrance (%)	Reference

NPcis, B6	Astrocytoma	Up to 70	(61,=20 62)
GFAP-H-Ras^V12	Astrocytoma	95	(17)=20
S100β-v-erbB	Oligodendroglioma	63	(79)=20
ND2:SmoA1	Medulloblastoma	48	(23)=20

It is highly likely that mouse models using = different driver=20 mutations leading to brain cancer will identify different genetic = risk=20 factors, since different tumor initiating events may be modified = by=20 different factors. It will therefore be important to interpret = findings in=20 mouse models according to subtypes of human brain tumors. Careful=20 characterization of mouse models of brain tumors will be = important, so=20 that findings can be validated in the most relevant subpopulation = of human=20 brain tumors. The Cancer Genome Atlas is characterizing human=20 glioblastomas at the molecular level and will help to better align = human=20 glioblastoma subpopulations with the closest available mouse = model. As an=20 example, recent studies have found that a subset of glioblastomas = carry=20 mutations in NF1(48,=20 57) and genetic risk loci identified in the NPcis=20 mouse model (61)=20 may be most relevant in patients with this subset of tumors. As = discussed=20 above, attempts to validate results from specific mouse models = across the=20 entire human population may fail because of the swamping out of = specific=20 subpopulation effects.

=20

=20

". . . A PIGMY AND A = PATAGONIAN,=20 A MOUSE AND A MAMMOTH . . ."

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Thomas Jefferson noted in his "Notes on the State = of=20 Virginia"(33)=20 that although animals of different shapes and sizes had much in = common,=20 ". . . all the manna in heaven would never raise = the Mouse=20 to the bulk of the Mammoth." Although there is much to learn from=20 comparisons of mouse models and human brain tumors to better = understand=20 risk factors, it is clear there will be species-specific risk = factors that=20 cannot be discovered in model systems. Due to differences in = metabolism,=20 mice may respond to dietary and environmental risk factors = differently=20 than humans. The differences in the size of the mouse and its = lifespan=20 mean that mouse brain tumors are in some ways intrinsically = different from=20 human brain tumors. Nonetheless, studies of cancer in mice have=20 demonstrated that many of the fundamental pathways are the same as = in=20 humans. It is important to note that mouse models provide a way of = getting=20 at some of the fundamental mechanisms underlying genetic risk = using a more=20 controlled experimental system. An understanding of these = mechanisms can=20 then guide studies in humans toward more focused questions. As an = example,=20 one may be able to define different genetic subgroups in the human = population and use these subgroups to then examine the effects of=20 environmental factors or diet. It will therefore be important to = approach=20 the understanding of risk for brain tumors from both human genome = wide=20 association studies, with the clear strengths of looking for = strong=20 effects on brain tumors in the human population, and from mouse = complex=20 trait studies, with the strengths of being able to isolate = different=20 variables and use a more controlled systems genetics approach to=20 understand more complex genetic interactions.

=20

=20

IMPLICATIONS FOR UNDERSTANDING RISK = FACTORS FOR=20 BRAIN TUMORS

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=

It has been recently recommended by the American = Academy of=20 Pediatrics that children between the ages of 2 and 10 be screened = for high=20 cholesterol, and that children with high cholesterol modify diet,=20 lifestyle or take medications to lower cholesterol and reduce the = risk of=20 cardiovascular disease (14).=20 This recommendation received a lot of media attention (56),=20 both positive and negative, and it highlights how in very common = diseases,=20 such as heart disease and diabetes, medical research has been able = to=20 identify risk factors and implement early screening and early = intervention=20 in patient populations. It remains to be seen over the next = generation how=20 effective these interventions are at lowering the incidence of = common=20 diseases in the population. The ability to screen and intervene = early=20 depends on an understanding of the mechanism underlying the = increased=20 risk. By studying brain tumor risk in both humans and mice, it is = hoped=20 that markers of increased risk can be identified to facilitate = early=20 screening and that the mechanism can be understood to allow for=20 intervention and reduction of risk.

=20

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=20

REFERENCES

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1.
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