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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_01C9B9FC.E0057140 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_01C9B9FC.E0057140 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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Journal of Neurochemistry

ISSN: 1471-4159 (Online)

ISSN: 0022-3042 (Print)

Volume 108, Issue 3, 2009.

Pages: 563=96577

Top of Page
Abstract
Materials 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 Journal compilation = © 2009 International Society for Neurochemistry

'); } =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 RT-PCR and western blot = analysis of sodium-ascorbic acid transporter 2. (a) The PCR products = were separated on 1.2% agarose gels and visualized by staining with = ethidium bromide. Upper panel: Lane 1, DNA 100-mer size standard. The = numbers on the left indicate the base pairs. Lane 2, positive controls = using mRNA isolated from fetal human brain. Lane 3, RT-PCR products = obtained using mRNA poly A+ isolated from 17-day-old embryonic rat brain = cortex. Lane 4, RT-PCR products obtained using mRNA poly A+ isolated = from cerebellar cortex. Lane 5, RT-PCR products obtained using mRNA = isolated from cultured embryonic rat brain cortex neurons. Lane 6, = reactions in which the cDNA synthesis step was performed in absence of = the reverse transcriptase (RT−) using the mRNA isolated from = cultured embryonic rat brain cortex neurons. Lower panel, RT-PCR for = β-actin was performed as an internal control. (b) Total protein = extracts were prepared from embryonic total brain extract (lane 1), = cultured 17-day-old embryonic rat brain cortex neurons (lane 2), = 17-day-old embryonic rat brain cortex tissue (lane 3), and human = embryonic kidney 293 cells (lane 4). A single band migrating with an = apparent molecular mass of 65 kDa was detected.'); }=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 In situ = hybridization and immunocytochemical characterization of SVCT2 = expression in embryonic rat brain. Left panel: SVCT2 mRNA expression in = 17-day-old embryonic mouse brain. Sagital brain sections hybridized = using a 620 bp digoxigenin-labeled riboprobes specific for SVCT2. = (a) Brain cortex observed with low magnification (lateral ventricle; = LV). Arrows indicate the positive reaction. (b) Control using sense = riboprobe (adjacent section to a). (c) Brain cortex observed with high = magnification. Arrows indicate the positive reaction (VZ, ventricular = zone; SVZ, subventricular zone). (d) Control using sense riboprobe = (adjacent section to c). (e) Ventricular wall of the lateral ventricle. = The arrows indicate a lower reaction present in the external area of the = cortex (cortical plate; CP). (f) Choroid plexus cells from lateral = ventricle. Inset, high magnification analysis of the epithelial cells = showing positive reaction (arrow). (g) Brain matrix from the lateral = ventricle. (h) Choroid plexus cells from fourth ventricle (IV–V). = The positive reaction was mainly observed in the SVZ of the brain cortex = (a–e), VZ of the brain matrix (g), and choroids plexus cells (f = and h). Right panel: (i–l) Immunohistochemical analysis of SVCT2 = expression. (l–k) Sections were incubated with anti-SVCT2 and the = immunoreaction was developed using the peroxidase anti-peroxidase = complex method. The positive reaction was detected in the = periventricular zone of the ventricular system (i–j) and in the = choroids plexus cells (k). (l) Control. CC, brain cortex; LV, lateral = ventricle; CP, choroidal plexus; H, hippocampus; M, marginal zone. Scale = bars: (a–b) 200 μm; (c–h) 50 μm; = (i–j) 100 μm; (k–l) 150 μm.'); }=0A= function display_f2(size) { var i =3D new Array(); i[0] =3D 'nf2'; i[1] = =3D 'nf2'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f2', size); imagelink(hwnd, i[size]); = caption_f2(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f3(hwnd) = {hwnd.document.write('Fig. 3 SVCT2 expression in the = rat embryonic brain using double-labeling analysis. Sagital sections of = embryonic brain were stained with anti-SVCT2 (b, e, and h) and a = secondary antibody Cy2-labeled (green). The nucleous of cells were = detected using propidium iodide (red, a, d, and g). The merged images of = both staining are shown in (c, f and i). (a–c) Brain cortex: = positive staining of cells localized in the ventricular and = subventricular zone (SVZ). Inset in (c). The migrating cortical = neuroblasts show the highest reaction. (d–f) Brain cerebellum: = positive reaction observed in the external granular cortex cells (eG) = and the neuroblasts of the internal granular cortex (arrow and inset in = f). (g–i) Choroid plexus: the propidium iodide staining was very = weak. The immunoreaction for anti-SVCT2 was observed in the cellular = membrane and the cytoplasm of the epithelial cells (arrow). Inset in = (i). Detailed analysis of the positive reaction detected in the = epithelial cells. LV, lateral ventricle. Scale bars: = 20 μm.'); }=0A= function display_f3(size) { var i =3D new Array(); i[0] =3D 'nf3'; i[1] = =3D 'nf3'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f3', size); imagelink(hwnd, i[size]); = caption_f3(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f4(hwnd) = {hwnd.document.write('Fig. 4 Immunohistochemical = characterization of SVCT2 expression and kinetic analysis of vitamin C = uptake in embryonic rat neurons isolated from brain cortex. Cultured = neurons obtained from 17-day-old rat embryos. Left panel: The = immunofluorescence analysis was performed using secondary antibodies = labeled with Cy2 (green). The nucleus was stained with propidium iodide = (red). (a) After 7 days in culture, cluster aggregation of the = cells was observed and most of the soma and neuronal processes showed an = intense immunoreaction with anti-MAP2. (b) Only 1% of the cultured cells = were positives with anti-GFAP (arrow). (c) The soma and cell processes = of the neurons also showed an intense staining with anti-SVCT2 (arrows). = (d) The neurons were negatives for anti-SVCT1. (e–h) Analysis of = vitamin C transport in primary cultures of embryonic rat neurons. (e) = Time-course uptake of 100 μM AA in the presence of NaCl or = replacing NaCl with choline chloride to 37°C. (f) Time-course uptake = of 100 μM AA in the presence of NaCl and cytochalasin B = (inhibitor of glucose and dehydroascorbic acid uptake) or replacing NaCl = with choline chloride and cytochalasin B. (g–h) Kinetic analysis = of the AA transport. Dose–response of the transport of AA to = 37°C (g). Eadie–Hofstee plot for the substrate dependence of = AA transport (h). Data represent the mean ± SD of three = experiments. Scale bars: (a–d) 15 μm.'); }=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 Immunohistochemical = characterization of SVCT2 expression and kinetic analysis of vitamin C = uptake in embryonic rat neurons isolated from cerebellar cortex. = Cultured cerebellar neurons obtained from 1-day-old postnatal rat. = (a–c) The immunostaining was developed using the peroxidase = anti-peroxidase complex method. (a) After 7-days in culture, cluster = aggregation of the cells was observed and most of the soma and neuronal = processes showed an intense immunoreaction with anti-neuronal specific = enolase. (b) The astrocytes were detected with anti-GFAP. The cells = showed a radial morphology and represented 20% of the cultured cells. = (c) The soma and cell processes of the neurons showed an intense = staining with anti-SVCT2. (d–e) Analysis of vitamin C transport = in primary cultures of cerebellar rat neurons. (d) Uptake of = 100 μM AA in the presence of NaCl or replacing NaCl with = choline chloride to 37°C. (e) Kinetic analysis of the AA transport. = Dose–response of the transport of AA to 37°C. (f) = Eadie–Hofstee plot for the substrate dependence of AA transport. = Data represent the mean ± SD of two experiments. Scale = bars: (a–c) 30 μm.'); }=0A= function display_f5(size) { var i =3D new Array(); i[0] =3D 'nf5'; i[1] = =3D 'nf5'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f5', size); imagelink(hwnd, i[size]); = caption_f5(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f6(hwnd) = {hwnd.document.write('Fig. 6 Immunohistochemical = characterization and RT-PCR analysis of SVCT2 expression in embryonic = rat neurons, Neuro2a, and HN33.11 cell lines. An intense immunoreaction = for tubulin and SVCT2 was detected in 7-day cultured neurons = (a–c), neuroblastoma Neuro2a (d–f) and HN33.11 = (g–i) cell lines. In neurons, the immunoreaction was concentrated = in the soma and cell processes. However, in immortalized cells the = reaction was mainly observed intracellularly. (j) RT-PCR analysis for = SVCT2 and β-actin expression. The PCR products were separated on = 1.2% agarose gels and visualized by staining with ethidium bromide. Lane = 1, DNA 100-mer size standard. The numbers on the left indicate the base = pairs. Lane 2, positive controls using mRNA isolated from brain cortex = neurons. Lane 3, RT-PCR products obtained using mRNA isolated from = Neuro2a cell lines. Lane 4, RT-PCR products obtained using mRNA isolated = from HN33.11 cell lines. Lane 5, reactions in which the cDNA synthesis = step was performed in absence of the reverse transcriptase (RT−) = using the mRNA isolated from brain cortex neurons. Scale bars: = (a–i) 12 μm.'); }=0A= function display_f6(size) { var i =3D new Array(); i[0] =3D 'nf6'; i[1] = =3D 'nf6'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f6', size); imagelink(hwnd, i[size]); = caption_f6(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f7(hwnd) = {hwnd.document.write('Fig. 7 Immunohistochemical = characterization of SVCTs and GLUT1 expression in human neuroblastoma = tumors. The immunofluorescence analysis was performed using secondary = antibodies labeled with Cy2 (green). The nucleus was stained with = propidium iodide (red). Intense immunoreaction for SVCT2 and GLUT1 was = detected in a neuroblastoma tumor. SVCT2 was detected in the cellular = membrane (a–c), however; GLUT1 immunoreaction was observed = intracelullarly (g–i). SVCT1 was not detected in neuroblastoma = tumor cells. Scale bars: 20 μm.'); }=0A= function display_f7(size) { var i =3D new Array(); i[0] =3D 'nf7'; i[1] = =3D 'nf7'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f7', size); imagelink(hwnd, i[size]); = caption_f7(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f8(hwnd) = {hwnd.document.write('Fig. 8 Kinetic analysis of = vitamin C uptake in neuroblastoma cell lines. Analysis of vitamin C = transport in Neuro2a and HN33.11 cell lines. (a and e) Time-course = uptake of 100 μM AA in the presence of NaCl or replacing = NaCl with choline chloride to 37°C.(b–f)) Time-course uptake = of 25, 50, and 100 μM AA in the presence of NaCl to = 37°C. (c and g) Kinetic analysis of the AA transport. = Dose–response of the transport of AA to 37°C. = Eadie–Hofstee plot for the substrate dependence of AA transport = (d and h). Data represent the mean ± SD of three = experiments.'); }=0A= function display_f8(size) { var i =3D new Array(); i[0] =3D 'nf8'; i[1] = =3D 'nf8'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f8', size); imagelink(hwnd, i[size]); = caption_f8(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= function caption_f9(hwnd) = {hwnd.document.write('Fig. 9 Acute effect of inhibitors = on the initial rates of AA uptake. Brain and cerebellar cortex neurons, = Neuro2a, and HN33.11 cells lines were pre-treated with the different = inhibitors for 10 min. Time-course uptake of 100 μM = ascorbic acid (AA) replacing NaCl with choline chloride (1), = 200 μM quercetin (2), 300 μM phloretin (3), = cytochalasin E (4), and cytochalasin B (5). Semi-log plot of the = concentration dependence for inhibition of AA transport by quercetin in = brain cortex neurons (C) or Neuro2a cell lines (F). Data represent the = mean ± SD of two experiments.'); }=0A= function display_f9(size) { var i =3D new Array(); i[0] =3D 'nf9'; i[1] = =3D 'nf9'; var hwnd =3D newfigwin(); dochead(hwnd, ''); copyright(hwnd); = otherlinks(hwnd, 'f9', size); imagelink(hwnd, i[size]); = caption_f9(hwnd); doctail(hwnd); hwnd.document.close(); hwnd.focus(); }=0A= =0A= =0A= function imagelist() {var hwnd =3D newfigwin(); dochead(hwnd, 'List of = Images - '); copyright(hwnd); =0A= hwnd.document.write('');=0A= =0A= =0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');=0A= hwnd.document.write('');hwnd.document.write('

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Journal of Neurochemistry

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Volume 108, Issue 3, Pages 563-577

Published Online: 19 Nov 2008

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The = Na⁺-dependent l-ascorbic acid transporter SVCT2 = expressed in=20 brainstem cells, neurons, and neuroblastoma cells is inhibited by=20 flavonoids

Teresa Caprile,=20 Katterine Salazar, Allisson Astuya,=20 Pedro Cisternas, Carmen Silva-Alvarez, Hern=E1n Montecinos, Carola Mill=E1n,=20 Mar=EDa de los = Angeles Garc=EDa and Francisco Nualart

Department of = Cellular Biology,=20 Faculty of Biological Sciences, University of Concepcion, = Concepcion,=20 Chile

Address correspondence and = reprint=20 requests to Dr Francisco Nualart, Departamento de Biolog=EDa = Celular,=20 Facultad de Ciencias Biol=F3gicas, Casilla 160-C, Universidad de = Concepci=F3n,=20 Concepci=F3n, Chile. E-mail: frnualart@udec.cl

Copyright Journal compilation =A9 2009 = International Society for Neurochemistry

KEYWORDS

brain cortex =95 flavonoid =95 neuron =95 = sodium=96vitamin C=20 co-transporters 2 =95 stem cells

ABSTRACT

3DAbstract=20

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Ascorbic acid (AA) is best known for its role as = an=20 essential nutrient in humans and other species. As the brain does = not=20 synthesize AA, high levels are achieved in this organ by specific = uptake=20 mechanisms, which concentrate AA from the bloodstream to the CSF = and from=20 the CSF to the intracellular compartment. Two different isoforms = of=20 sodium=96vitamin C co-transporters (SVCT1 and SVCT2) have been = cloned. Both=20 SVCT proteins mediate high affinity = Na⁺-dependent=20 l-AA transport and are = necessary for the=20 uptake of vitamin C in many tissues. In the adult brain the = expression of=20 SVCT2 was observed in the hippocampus and cortical neurons by = in situ hybridization; however, there is no data = regarding=20 the expression and distribution of this transporter in the fetal = brain.=20 The expression of SVCT2 in embryonal mesencephalic neurons has = been shown=20 by RT-PCR suggesting an important role for vitamin C in = dopaminergic=20 neuronal differentiation. We analyze SVCT2 expression in human and = rat=20 developing brain by RT-PCR. Additionally, we study the normal = localization=20 of SVCT2 in rat fetal brain by immunohistochemistry and in=20 situ hybridization demonstrating that SVCT2 is highly = expressed in=20 the ventricular and subventricular area of the rat brain. SVCT2 = expression=20 and function was also confirmed in neurons isolated from brain = cortex and=20 cerebellum. The kinetic parameters associated with the transport = of AA in=20 cultured neurons and neuroblastoma cell lines were also studied. = We=20 demonstrate two different affinity transport components for AA in = these=20 cells. Finally, we show the ability of different flavonoids to = inhibit AA=20 uptake in normal or immortalized neurons. Our data demonstrates = that brain=20 cortex and cerebellar stem cells, neurons and neuroblastoma cells = express=20 SVCT2. Dose-dependent inhibition analysis showed that quercetin = inhibited=20 AA transport in cortical neurons and Neuro2a = cells.

Received October 3, 2008; accepted October 30,=20 2008.

DIGITAL OBJECT IDENTIFIER (DOI)

10.1111/j.1471-4159.2008.05788.x About=20 DOI

=20

=20

Article Text

Two mechanisms of vitamin C transport have been = suggested.=20 One involves the efficient transport of the oxidized form of = vitamin C,=20 dehydroascorbic acid (DHA), through glucose transporters (GLUTs) = (Agus=20 et al. 1999; Nualart=20 et al. 2003), and the other = involves the=20 transport of the reduced form of vitamin C, ascorbic acid (AA), = through=20 sodium=96vitamin C transporters (SVCT1 and SVCT2) (Faaland=20 et al. 1998; Daruwala=20 et al. 1999; Rajan=20 et al. 1999; Tsukaguchi=20 et al. 1999; Wang=20 et al. 1999, 2000). Functional = data suggests=20 a novel role for AA in dopamine neuronal differentiation, which is = independent of its antioxidative properties (Yan=20 et al. 2001). We have demonstrated = the=20 expression of the SVCT2 transporter mRNA in mouse brain cortical = area (Castro=20 et al. 2001). Furthermore, our = kinetic=20 analysis and the inhibition data have indicated the expression of = SVCT2=20 transporter in primary cultured mouse cortical neurons (Castro=20 et al. 2001, 2007). Similar = results were=20 recently demonstrated by Qiu=20 et al. (2007) using cortical = neurons=20 prepared from wild-type Slc23a+/+ or Slc23a−/− mice. = In adult brain, SVCT2=20 is expressed in neurons, microglia, and ependymal cells (Berger=20 and Hediger 2000;Korkok=20 et al. 2000; Castro=20 et al. 2001; Garc=EDa=20 et al. 2003; Lee=20 et al. 2003; Astuya=20 et al. 2005; Mun=20 et al. 2006). However, = immunohistochemical=20 analyses have not been performed to determine the exact = distribution of=20 this transporter in the embryonic brain.

Vitamin C levels are particularly high in fetal = rat brain,=20 doubling from the 15th to the 20th day of gestation and then = dropping=20 significantly by the time of birth (Kratzing=20 et al. 1985). This suggests a role = of=20 vitamin C during brain development. However, apparent normal brain = differentiation has been observed in SVCT2-null mice, suggesting = that=20 embryonic cells may use different mechanisms or transporters to = take up=20 vitamin C (Sotiriou=20 et al. 2002). SVCT2 mRNA has been = detected=20 in cultured dopaminergic neurons, cultured brain cortex mouse = neurons,=20 isolated brain cortical precursor and neuroblastoma cells (Castro=20 et al. 2001; Yan=20 et al. 2001; Lee=20 et al. 2003; Astuya=20 et al. 2005). Additional data = indicate that=20 SVCT2 is also expressed in the hippocampal neuron (Qiu=20 et al. 2007). Previous data have = also shown=20 that neuronal hippocampal cultures from SVCT2-deficient mice have=20 increased susceptibility to oxidative damage; thus, it has been = proposed=20 that SVCT2 activity is needed for neuronal development, functional = maturation, and antioxidant responses (Qiu=20 et al. 2007). However, the = presence of a=20 given mRNA is not always synonymous with protein expression and = functional=20 activity; therefore, it is necessary to analyze both the mRNA = expression=20 and the protein distribution to obtain more definitive information = regarding the expression and localization of SVCT2 in fetal brain = as well=20 as the general function of vitamin C during brain development.

Herein, we demonstrate that SVCT2 transporter is = highly=20 expressed in the cortex and cerebellum of rat fetal brain. Kinetic = analysis of the AA uptake demonstrated the functional expression = of=20 SVCT2-like proteins in neurons isolated from brain cortex and = cerebellum.=20 In both types of neurons we observed the presence of two affinity=20 constants for the uptake of AA. A similar result was obtained = using=20 neuroblastoma cells.

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Materials and methods

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Tissue immunocytochemistry

Rat (Holtzmann) total brain extracts of embryonic = 17-day old=20 was dissected and fixed by immersion in Bouin's solution. Human=20 neuroblastoma biopsies were also fixed in Bouin's solution. = Samples were=20 dehydrated in graded alcohol solutions and embedded in paraffin. = Frontal=20 and sagital sections (7 μm) of rat brain and tumor = samples were=20 mounted on poly-l-lysine-coated = glass=20 slides. Endogenous peroxidase activity of xylene deparaffinized, = and=20 rehydrated sections were inhibited by treatment with 0.3%=20 H₂O₂/methanol. For = immunohistochemical analyses, we used an affinity-purified goat = polyclonal=20 antibody raised against a peptide mapping within the extracellular = domain=20 of SVCT2 of rat origin (Santa Cruz Biotechnology, Santa Cruz, CA, = USA).=20 The affinity purified goat polyclonal anti-SVCTs antibodies = diluted=20 1 : 100 were incubated overnight at 22=B0C. The = antibodies were=20 diluted in incubation buffer Tris=96HCl (pH 7.8) containing = 8.4 mM=20 sodium phosphate, 3.5 mM potassium phosphate, 120 mM = NaCl, and=20 1% bovine serum albumin (Nualart=20 et al. 1999). After extensive = washing,=20 sections were incubated for 2 h at 22=B0C with anti-goat IgG = peroxidase=20 labeled (1 : 50; Dako, Carpinteria, CA, USA). The = peroxidase=20 activity was developed using diaminobenzidine and=20 H₂O₂. As controls, = we=20 utilized primary antibodies pre-absorbed with pre-immune serum and = the=20 respective peptides used to elicit them. Slides were dehydrated = through=20 graded alcohol to xylene washing and mounted on coverslips. = Hematoxylin=20 was used for nuclear counterstaining. For confocal microscopy, = sections=20 were incubated for 2 h with Cy2 conjugated affinity purified = donkey=20 anti-goat IgG (1 : 200; Jackson ImmunoResearch, West = Grove, PA,=20 USA) at 22=B0C. Additionally, the tissue sections were incubated = with=20 propidium iodide after Rnase treatment for nuclear staining (Garc=EDa=20 et al. 2003).

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In situ = hybridization

A cDNA of approximately 0.6 kb subcloned in=20 pCR-4-Blunt-Topo (Clontech, Palo Alto, CA, USA) and encoding the = rat SVCT2=20 was used to generate sense and antisense digoxigenin-labeled = riboprobes.=20 RNA probes were labeled with digoxigenin-UTP by in = vitro transcription with T3 or T7 RNA polymerase following = the=20 manufacturer's procedure (Boehringer Mannheim, Mannheim, Germany). = In situ hybridization was performed on fetal = brain sections=20 mounted on poly-l-lysine-coated = glass=20 slides. The sections were deparaffinized in xylene and rehydrated = in=20 graded ethanol. Following proteinase K treatment [3 min at = 37=B0C in=20 phosphate-buffered saline (PBS), 1 μg/mL proteinase K], = the tissue=20 sections were fixed with 4%p-formaldehyde = for=20 5 min at 4=B0C, washed in cold PBS and then acetylated in = 0.1 M=20 triethanol amine=96HCl (pH 8.0) at 22=B0C for 10 min. After a = brief wash,=20 the sections were incubated in pre-hybridization solution for = 15 min=20 at 37=B0C, and then 25 μL of hybridization mix (50% = formamide,=20 0.6 M NaCl, 10 mM Tris=96HCl, pH 7.5, 1 mM EDTA, 1x = Denhart's=20 solution, 10% polyethilenglycol 8000, 10 mM dithiothreitol,=20 500 μg yeast tRNA/mL, 50 μg/mL heparin, = 500 μg/mL DNA=20 carrier, and 1 : 20 to 1 : 100 diluted = riboprobe) were=20 added to each slide. The slides were covered with glass coverslips = and=20 placed in a humidified chamber at 42=B0C overnight. After removal = of the=20 coverslip, slides were rinsed in 4x sodium citrate solution and = washed=20 twice for 15 min at 42=B0C. The slides were washed at 37=B0C = for=20 15 min each in 2x, 1x, and 0.3x sodium citrate solution.=20 Visualization of digoxigenin was performed by incubation with a = monoclonal=20 antibody coupled to alkaline phosphatase = (anti-digoxigenin-alkaline=20 phosphatase Fab fragments diluted 1 : 500; Boehringer = Mannheim)=20 for 2 h at 22=B0C. Nitroblue tetrazolium chloride and=20 5-bromo-4-chloro-3-indolyl-phosphate (Boehringer Mannheim) were = used as=20 substrates for the alkaline phosphatase. Controls included use of = the=20 sense riboprobe and omission of the probe as previously specified = (Garc=EDa=20 et al. 2001).

=20

=20

Cell culture and = immunocytochemistry

=20

=20

Primary cultures of neurons

Cultured neurons were obtained from Holtzmann = Rats.=20 Forebrain was isolated from 17-day-old embryos, while the = cerebellum was=20 obtained from 1-day postnatal rats. Tissues were digested for = 15 min=20 with trypsin 0.12% (wt/vol) in phosphate buffer 0.1 M (pH = 7.4,=20 320 mOsm) and triturated to homogeneity with a fire-polished = Pasteur=20 pipet. The cells were plated at 500 000 cells/mL onto = glass=20 coverslips coated with poly-l-lysine=20 (mol. Wt > 350 kDa; Sigma, St Louis, MO, USA) in = Dulbecco's modified Eagle's medium supplemented with 10% fetal = bovine=20 serum. After 30 min, the floating cells were removed and the = attached=20 cells were incubated and fed every 3 days with Neurobasal/B27 = medium=20 (Gibco BRL, Grand Island, NY, USA). The cultures were incubated in = 5%=20 CO₂ in a humidified environment at=20 37=B0C.

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Cell line culture

The mouse neuroblastoma cell line Neuro-2a and the = hippocampal=96neuroblastoma hybrid cell line HN33.11 (a gift of Dr = Marshall=20 Horwitz from Albert Einstein College of Medicine), established by = fusion=20 of mouse hippocampal cells with neuroblastoma cells (N18TG2), were = grown=20 in Dulbecco's modified Eagle's medium supplemented with 10% fetal = bovine=20 serum, penicillin, and streptomycin (Jin=20 et al. 1998; Evangelopoulos=20 et al. 2004).

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Immunocytochemistry

Neurons and neuroblastoma cells were grown on = coverslips,=20 fixed with 4%p-formaldehyde in PBS for = 30 min at=20 22=B0C, washed with PBS and incubated in PBS containing 1% bovine = serum=20 albumin and 0.2% Triton X-100 for 5 min at 22=B0C. The cells = were=20 incubated overnight at 22=B0C with the following antibodies:=20 anti-neuron-specific enolase (1 : 200; Dako),=20 anti-microtule-associated protein 2 (1 : 200; Boehringer = Mannheim), anti-glial fibrilary acidic protein (1 : 500; = Dako),=20 anti-tubulin βIII (1 : 2000, clone Tuj1; Promega, = Madison, WI,=20 USA) or anti-SVCT2 (1 : 100; Santa Cruz Biotechnology). = Cells=20 were incubated with goat anti-rabbit IgG or rabbit anti-mouse IgG=20 (1 : 100; Dako) for 2 h and them for 30 min = with=20 peroxidase anti-peroxidase complex (1 : 100; Dako). The=20 enzymatic activity was developed using diaminobenzidine and = hydrogen=20 peroxide (Nualart=20 et al. 1999). Alternatively, the = cultured=20 cells were incubated for 2 h with Cy2 conjugated affinity = purified=20 donkey anti-goat IgG, donkey anti-rabbit IgG, or donkey anti-mouse = IgG=20 (1 : 200; Jackson ImmunoResearch) at 22=B0C. The cells = were=20 incubated with propidium iodide after Rnase treatment for nuclear=20 staining. Negative controls for SVCT1 and SVCT2 consisted of = pre-immune=20 serum and both primary antibodies pre-absorbed with the respective = peptides used to raise them.

=20

=20

Immunoblotting

For immunoblot analysis, total protein from = embryonic brain=20 and brain cortex (17-day-old embryos), rat cultured neurons and = human=20 embryonic kidney 293 cells was obtained by homogenizing the cells = in=20 0.3 mM sucrose, 3 mM dithiothreitol, 1 mM EDTA,=20 100 μg/mL phenylmethylsulfonyl fluoride, = 1 μg/mL pepstatin,=20 2 μg/mL aprotinin, and 2 μg/mL leupeptine. = Total membrane=20 fractions were collected by high-speed centrifugation=20 (122 000 g, 60 min). Thirty = microgram=20 of protein was loaded in each lane, separated by polyacrylamide = gel=20 electrophoresis in the presence of sodium dodecyl sulfate, = transferred to=20 nitrocellulose membranes, and probed with anti-SVCT2 or = pre-absorbed=20 antibodies (1 : 500=961 : 1500) (Garc=EDa=20 et al. 2003). The secondary = antibody was=20 rabbit anti-goat IgG coupled to peroxidase (1 : 5000). = The=20 reaction was developed with enhanced chemiluminescence using the = ECL=20 western blotting analysis system (Amersham Corporation, Arlington = Heights,=20 IL, USA).

=20

=20

RT-PCR

The following samples were used for RT-PCR = analysis: (i)=20 embryonic rat brain cortex tissue, (ii) embryonic rat cerebellar = cortex=20 tissue, (iii) 7-day cultured neurons isolated from 17-day-old = embryos of=20 Holtzmann rats, (iv) Fetal mRNA isolated from human brain = (Clontech), and=20 (v) mRNA isolated from Neuro2a and HN33.11 cell lines. The poly(A) = RNA of=20 the neuron and brain tissues was isolated using the Oligotex = direct kit=20 (Qiagen, Valencia, CA, USA). For RT-PCR, 0.5=961 μg of = RNA was=20 incubated in 20 μL reaction volume containing 10 mM = Tris, pH=20 8.3, 50 mM KCl, 5 mM MgCl₂, = Rnase=20 inhibitor 20 U, 1 mM dNTPs, 2.5 μM of random=20 hexanucleotides, and 50 U of MuLV reverse transcriptase = (Perkin=20 Elmer, Waltham, MA, USA) for 10 min at 23=B0C followed by = 30 min=20 at 42=B0C and 5 min at 94=B0C. Parallel reactions were = performed in the=20 absence of reverse transcriptase to control for the presence of=20 contaminant DNA. For amplification, a cDNA aliquot in a volume of=20 12 μL containing 60 mM Tris, pH 9.1, 18 mM=20 = (NH₄)₂SO₄,=20 7 mM MgCl2, 0.2 mM dNTPs, = 0.04 U of=20 elongase (Gibco BRL), and 0.4 μM primers was incubated = at 94=B0C for=20 4 min, 94=B0C for 20 s, 54=B0C for 30 s, and 68=B0C = for 40 s=20 for 35 cycles. PCR products were separated by 1.2% agarose gel=20 electrophoresis and visualized by staining with ethidium bromide. = The=20 following primers (based on human sequence F164142) were used to = analyze=20 the expression of SVCT2 transporter: forward primer=20 5'-TTGACCATTACACCCACGGT-3' and reverse primer = 5'-GTAATTCCCAAAACTCCAAT-3'=20 (expected product 620 bp) or forward primer=20 5'-TTCTGTGTGGGAATCACTAC-3' and reverse primer = 5'-ACCAGAGAGGCCAATTAGGG-3'=20 (expected product 339 bp).

=20

=20

Vitamin C uptake analysis

After 5 days in culture the neurons were used = for=20 kinetic analysis. Cultures were carefully selected under the = microscope to=20 ensure that only plates showing uniformly growing cells were used. = In each=20 experiment, cells from 12 wells incubated with buffer were removed = and=20 used to estimate the average total cell number present in each = well. We=20 did not observe a great variation in cell number between the wells = after=20 buffer incubation (Nualart=20 et al. 2003). The cells were = washed with=20 incubation buffer (15 mM HEPES, 135 mM NaCl, 5 mM = KCl,=20 1.8 mM CaCl₂, and 0.8 mM=20 MgCl₂) and incubated in the same medium = for=20 5 min at 22=B0C. A similar protocol was used with Neuro2a and = HN33.11=20 cell lines. Uptake assays were performed in 500 μL of = incubation=20 buffer containing 0.1 μCi of = 1-¹⁴C-l-AA (specific activity = 4 mCi/mmol;=20 DuPont-NEN, Boston, MA, USA), to a final concentration of = 1=96300 μM.=20 The uptake was stopped by washing the cells with ice-cold PBS. = Cells were=20 dissolved in 0.5 mL of lysis buffer (10 mM Tris=96HCl, = pH 8.0, and=20 0.2% sodium dodecyl sulfate), and the incorporated radioactivity = was=20 assayed by liquid scintillation spectrometry. The Michaelis = constant,=20 K_m, was calculated = using the=20 Eadie=96Hofstee analysis. Data represent mean =B1 SD of = three=20 experiments with each determination performed in triplicate. In = inhibition=20 experiments statistical comparison between two or more groups of = data was=20 carried out using anova = (followed by=20 Bonferroni post-test). A value of p < 0.05 was considered to be = statistically=20 significant.

=20

=20

Results

=

=

=

=

=20

=20

Rat and human embryonic brain = express=20 SVCT2

The expression of a sodium=96AA co-transporter in = rat=20 embryonic brain tissue was analyzed by RT-PCR with specific = primers for=20 the SVCT2 isoform. The conditions were optimized using RNA from = human=20 brain tissue as a control for the expression. As judged by their=20 migration, the amplified DNA band was approximately 620 bp, = the=20 expected size for the amplification products of SVCT2 (Fig. 1a,=20 lane 2). No amplification product was observed in samples in which = the=20 cDNA synthesis step was performed in the absence of reverse = transcriptase,=20 indicating the absence of DNA contamination in the RNA preparation = (Fig. 1a,=20 lane 6).

=20

= Fig. 1 = RT-PCR=20 and western blot analysis of sodium-ascorbic acid = transporter 2. (a)=20 The PCR products were separated on 1.2% agarose gels and = visualized=20 by staining with ethidium bromide. Upper panel: Lane 1, DNA = 100-mer=20 size standard. The numbers on the left indicate the base = pairs. Lane=20 2, positive controls using mRNA isolated from fetal human = brain.=20 Lane 3, RT-PCR products obtained using mRNA poly A+ isolated = from=20 17-day-old embryonic rat brain cortex. Lane 4, RT-PCR = products=20 obtained using mRNA poly A+ isolated from cerebellar cortex. = Lane 5,=20 RT-PCR products obtained using mRNA isolated from cultured = embryonic=20 rat brain cortex neurons. Lane 6, reactions in which the = cDNA=20 synthesis step was performed in absence of the reverse = transcriptase=20 (RT−) using the mRNA isolated from cultured embryonic = rat brain=20 cortex neurons. Lower panel, RT-PCR for β-actin was = performed as an=20 internal control. (b) Total protein extracts were prepared = from=20 embryonic total brain extract (lane 1), cultured 17-day-old=20 embryonic rat brain cortex neurons (lane 2), 17-day-old = embryonic=20 rat brain cortex tissue (lane 3), and human embryonic kidney = 293=20 cells (lane 4). A single band migrating with an apparent = molecular=20 mass of 65 kDa was detected.

[Normal=20 View ]

Using mRNA isolated from 17-day-old embryonic rat, = we=20 amplified the SVCT2 isoform from the brain cortex (Fig. 1a,=20 lane 3). Additionally, we detected SVCT2 expression in mRNA = isolated from=20 embryonic rat cerebellar cortex tissue (Fig. 1a,=20 lane 4). SVCT2 expression was also observed in 7-day cultured = neurons=20 isolated from brain cortex of 17-day-old rat embryos (Fig. 1a,=20 lane 5). The expression and concentration of the β-actin gene = in reactions=20 indicates the high quality of the RNAs used to perform the = analysis (Fig. 1b).=20 Our results indicate the early expression of SVCT2 transporters in = embryonic rat and human brain tissue. SVCT2 expression was also = evaluated=20 by western blotting using protein extracts isolated from cultured=20 17-day-old embryonic rat brain cortex neurons. Our analyses = indicate that=20 the neurons express a protein band of approximately 60 kDa = reactive=20 to anti-SVCT2 (Fig. 1b,=20 lane 2) that co-migrated with a similar protein band present in = extracts=20 of embryonic total brain, 17-day-old rat brain cortex tissue (Fig. 1b,=20 lanes 1, 3) and human embryonic kidney 293 cells (Fig. 1,=20 lane 4). No immunoreactive proteins were detected when anti-SVCT2 = antibody=20 was pre-absorbed with the blocking peptide (data not = shown).

=20

=20

SVCT2 is highly expressed in = neurogenic=20 zones

Different studies have suggested that the = concentration of=20 AA is very high in fetal rat brain (Kratzing=20 et al. 1985); however, no = localization=20 analyses of the expression of AA transporters involved in the = uptake of=20 vitamin C in the different cells of the fetal brain have been = published.=20 We analyzed the gene expression of SVCT2 at the mRNA level by = in situ hybridization using digoxigenin-labeled = cRNA probes=20 specific for SVCT2. The positive hybridization signal was = concentrated in=20 the brain cortex facing the lateral ventricle (Fig. 2a,=20 arrows). The reaction observed in this area was completely = abolished when=20 the sense probe was used (Fig. 2b=20 and d). A detailed analysis of the reaction showed that the = highest=20 hybridization signal was detected in the subventricular zone (Fig. 2c,=20 SVZ) and in different migrating cells throughout the brain cortex = (Fig. 2c,=20 arrows). In specific areas of the brain a weak hybridization = signal was=20 detected in the external area of the brain cortex (Fig. 2e,=20 arrows). In this area, the localization of differentiated neurons = has been=20 demonstrated (Dehay=20 and Kennedy 2007). A high hybridization signal was also = detected in=20 the choroid plexus of the lateral ventricle (Fig. 2f=20 and inset), however, the choroid plexus of the fourth ventricle = showed=20 weak staining (Fig. 2h).=20 A detailed analysis of the different areas of the brain led us to = observe=20 a clear expression of SVCT2 mRNA in neuroepithelial cells of the=20 ventricular matrix (Fig. 2g,=20 VZ). To compare the SVCT2 mRNA distribution and the protein = localization=20 we used immunohistochemical analysis of SVCT2 transporter. The = positive=20 reaction was very intense in the neuroepithelial cells localized = around=20 the ventricular system (Fig. 2i,=20 arrows). The cells from the choroid plexus showed a weak = immunostaining in=20 the third ventricle [Fig. 2i,=20 cortical plate (CP)]. Similar pattern of immunostaining was = observed in=20 choroid plexus differentiated cells localized in the lateral = ventricles=20 (Fig. 2j=96k,=20 CP). However, the cells localized in the base of the choroid = plexus showed=20 intense immunostaining (Fig. 2k,=20 arrows and inset). In the cortex the immunostaining was localized = in the=20 VZ and the SVZ (Fig. 2i=96k).=20 A similar reaction was observed in migrating neurons throughout = the=20 hippocampus (Fig. 2j).=20 The external area of the brain cortex was almost negative; however = a weak=20 immunoreaction was detected in the CP near the marginal zone (Fig. 2i=96j).=20 No immunostaining was detected when the anti-SVCT2 antibody was=20 pre-absorbed with the blocking peptide (Fig. 2l).

=20

= Fig. 2 = In situ hybridization and = immunocytochemical=20 characterization of SVCT2 expression in embryonic rat brain. = Left=20 panel: SVCT2 mRNA expression in 17-day-old embryonic mouse = brain.=20 Sagital brain sections hybridized using a 620 bp=20 digoxigenin-labeled riboprobes specific for SVCT2. (a) Brain = cortex=20 observed with low magnification (lateral ventricle; LV). = Arrows=20 indicate the positive reaction. (b) Control using sense = riboprobe=20 (adjacent section to a). (c) Brain cortex observed with high = magnification. Arrows indicate the positive reaction (VZ,=20 ventricular zone; SVZ, subventricular zone). (d) Control = using sense=20 riboprobe (adjacent section to c). (e) Ventricular wall of = the=20 lateral ventricle. The arrows indicate a lower reaction = present in=20 the external area of the cortex (cortical plate; CP). (f) = Choroid=20 plexus cells from lateral ventricle. Inset, high = magnification=20 analysis of the epithelial cells showing positive reaction = (arrow).=20 (g) Brain matrix from the lateral ventricle. (h) Choroid = plexus=20 cells from fourth ventricle (IV=96V). The positive reaction = was mainly=20 observed in the SVZ of the brain cortex (a=96e), VZ of the = brain=20 matrix (g), and choroids plexus cells (f and h). Right = panel: (i=96l)=20 Immunohistochemical analysis of SVCT2 expression. (l=96k) = Sections=20 were incubated with anti-SVCT2 and the immunoreaction was = developed=20 using the peroxidase anti-peroxidase complex method. The = positive=20 reaction was detected in the periventricular zone of the = ventricular=20 system (i=96j) and in the choroids plexus cells (k). (l) = Control. CC,=20 brain cortex; LV, lateral ventricle; CP, choroidal plexus; = H,=20 hippocampus; M, marginal zone. Scale bars: (a=96b) = 200 μm; (c=96h)=20 50 μm; (i=96j) 100 μm; (k=96l) = 150 μm.=20

[Normal View=20 ]

To obtain more information about the specific = localization=20 of SVCT2 transporter in brain neurons we performed = immunofluorescence=20 analysis and confocal microscopy. The nuclei of the cells present = in the=20 brain cortex and cerebellar cortex were detected with propidium = iodide (Fig. 3).=20 SVCT2 expression was mainly observed in the periventricular region = of the=20 brain cortex and granular external layer of the cerebellum (Fig. 3b=96c=20 and e=96f). In these areas are mainly located proliferating = precursor=20 cells (Okano-Uchida=20 et al. 2004; Dehay=20 and Kennedy 2007). Similar immunostaining was observed in = migrating=20 cells (Fig. 3c=20 and f, insets). Using confocal microscopy and = immunofluorescence=20 analysis it was possible to detect a clear immunostaining in = choroid=20 plexus cells (Fig. 3h=96i).

=20

= Fig. 3 = SVCT2=20 expression in the rat embryonic brain using double-labeling=20 analysis. Sagital sections of embryonic brain were stained = with=20 anti-SVCT2 (b, e, and h) and a secondary antibody = Cy2-labeled=20 (green). The nucleous of cells were detected using propidium = iodide=20 (red, a, d, and g). The merged images of both staining are = shown in=20 (c, f and i). (a=96c) Brain cortex: positive staining of = cells=20 localized in the ventricular and subventricular zone (SVZ). = Inset in=20 (c). The migrating cortical neuroblasts show the highest = reaction.=20 (d=96f) Brain cerebellum: positive reaction observed in the = external=20 granular cortex cells (eG) and the neuroblasts of the = internal=20 granular cortex (arrow and inset in f). (g=96i) Choroid = plexus: the=20 propidium iodide staining was very weak. The immunoreaction = for=20 anti-SVCT2 was observed in the cellular membrane and the = cytoplasm=20 of the epithelial cells (arrow). Inset in (i). Detailed = analysis of=20 the positive reaction detected in the epithelial cells. LV, = lateral=20 ventricle. Scale bars: 20 μm. =

[Normal=20 View ]

=20

=20

SVCT2 is functionally expressed in = neurons=20 isolated from brain cortex and cerebellum

Although it is known that AA is present both in = neurons and=20 glial cells, transport systems in neurons have not been thoroughly = characterized. To demonstrate the neuronal functional expression = of SVCT2,=20 we isolated neurons from embryonic rat brain cortex. The cells = were=20 cultured for 7 days and the purity of the culture was = analyzed by=20 phase contrast microscopy and immunocytochemistry (Fig. 4a=20 and b and data not shown). The majority of the cells in = culture were=20 aggregated, forming clusters of different sizes, a typical = characteristic=20 of cultures rich in neurons (Fig. 4a,=20 arrows). Most of the cells in culture were positive for anti-MAP2=20 antiserum, a specific marker of neurons (Fig. 4a).=20 Only a few cells were positive to anti-GFAP, an astrocyte marker = (Fig. 4b,=20 arrow). In primary cultured neurons, we analyzed the expression of = SVCT=20 isoform by immunohistochemistry using the specific antibody to = SVCT2.=20 Intense immunoreaction was detected in the soma and processes of = the=20 neurons; however, the most intense reaction was observed around = the=20 nuclear area (Fig. 4c,=20 arrows). We were unable to observe any positive reaction = associated to=20 anti-SVCT1 specific antisera (Fig. 4d)=20 indicating that SVCT2 is selectively expressed in neurons.

=20

= Fig. 4=20 Immunohistochemical characterization of SVCT2=20 expression and kinetic analysis of vitamin C uptake in = embryonic rat=20 neurons isolated from brain cortex. Cultured neurons = obtained from=20 17-day-old rat embryos. Left panel: The immunofluorescence = analysis=20 was performed using secondary antibodies labeled with Cy2 = (green).=20 The nucleus was stained with propidium iodide (red). (a) = After=20 7 days in culture, cluster aggregation of the cells was = observed and most of the soma and neuronal processes showed = an=20 intense immunoreaction with anti-MAP2. (b) Only 1% of the = cultured=20 cells were positives with anti-GFAP (arrow). (c) The soma = and cell=20 processes of the neurons also showed an intense staining = with=20 anti-SVCT2 (arrows). (d) The neurons were negatives for = anti-SVCT1.=20 (e=96h) Analysis of vitamin C transport in primary cultures = of=20 embryonic rat neurons. (e) Time-course uptake of = 100 μM AA in=20 the presence of NaCl or replacing NaCl with choline chloride = to=20 37=B0C. (f) Time-course uptake of 100 μM AA in the = presence of=20 NaCl and cytochalasin B (inhibitor of glucose and = dehydroascorbic=20 acid uptake) or replacing NaCl with choline chloride and=20 cytochalasin B. (g=96h) Kinetic analysis of the AA = transport.=20 Dose=96response of the transport of AA to 37=B0C (g). = Eadie=96Hofstee plot=20 for the substrate dependence of AA transport (h). Data = represent the=20 mean =B1 SD of three experiments. Scale bars: = (a=96d)=20 15 μm.

[Normal View=20 ]

Next, we studied AA transport in rat neurons = isolated from=20 rat brain cortex and cerebellum. In rat brain cortex neurons, the=20 transport of AA was linear for the first 10 min (Fig. 4e).=20 The initial velocity of the AA uptake was=20 = 11 pmol =D7 10⁶cells/min. = The=20 total uptake in 10 min was=20 102 pmol =D7 10⁶ cells (Fig. 4e).=20 To test the sodium dependence of AA transport we replaced the NaCl = in the=20 incubation buffer with choline chloride. Transport of AA by = neurons was=20 decreased by at least 90% in the absence of sodium and the uptake = reached=20 a plateau of = 11 pmol =D7 10⁶ cells in=20 5 min (Fig. 4e).

It has been previously demonstrated that the = facilitative=20 GLUTs are efficient transporters of DHA, the oxidized form of = vitamin C=20 (Vera=20 et al. 1993; Rumsey=20 et al. 1997). Additionally, it has = also been=20 observed that neurons may take up DHA independent of sodium (Castro=20 et al. 2001). Because vitamin C = may be=20 spontaneously oxidized in vitro, we = analyzed the=20 effect of cytochalasin B, a GLUT inhibitor, in the uptake of AA, = the=20 reduced form of vitamin C (Fig. 4f).=20 As we observed previously, the transport of AA co-incubated with=20 cytochalasin B and sodium was linear for the first 10 min (Fig. 4f).=20 The initial velocity of AA uptake was=20 = 9 pmol =D7 10⁶cells/min. = The=20 total uptake in 10 min was=20 81 pmol =D7 10⁶ cells (Fig. 4f).=20 Transport of AA in neurons co-incubated with cytochalasin B and = choline=20 was decreased by at least 80% in the absence of sodium and the = uptake=20 reached a plateau of = 10 pmol =D7 10⁶=20 cells in 5 min (Fig. 4f).=20 These results strongly suggest the participation of a sodium=20 co-transporter in the transport of AA by neurons. We then = performed a=20 detailed kinetic analysis of the AA transport in neurons. = Dose=96response=20 studies using 5 min assays revealed that the transport of AA = in=20 neurons is saturated at concentrations higher than = 100 μM (Fig. 4g).=20 The Eadie=96Hofstee transformation showed apparent K_m of 11 and = 92 μM and V_max of 39 and=20 120 pmol =D7 10⁶/min, = respectively=20 (Fig. 4h).

To compare the functional expression of SVCT2 in = different=20 areas of the brain we examined the AA transporter present in rat = neurons=20 isolated from cerebellar cortex. The cells were cultured for = 7 days=20 and the purity of the culture was analyzed by phase contrast = microscopy=20 and immunocytochemistry (Fig. 5a=20 and b and data not shown). Most of the cells isolated from = cerebellar=20 cortex were aggregated forming clusters of different sizes (Fig. 5a,=20 arrow). The neurons in culture were positive for anti-neuronal = specific=20 enolase antiserum (Fig. 5a).=20 Only a few cells were positive for anti-GFAP, an astrocyte marker = (Fig. 5b,=20 arrow). Intense immunoreaction was detected using anti-SVCT2 = antibody and=20 localized in the soma and processes of the neurons (Fig. 5c,=20 arrows). We were unable to observe any positive reaction = associated to=20 anti-SVCT1 antisera (data not shown).

=20

= Fig. 5=20 Immunohistochemical characterization of SVCT2=20 expression and kinetic analysis of vitamin C uptake in = embryonic rat=20 neurons isolated from cerebellar cortex. Cultured cerebellar = neurons=20 obtained from 1-day-old postnatal rat. (a=96c) The = immunostaining was=20 developed using the peroxidase anti-peroxidase complex = method. (a)=20 After 7-days in culture, cluster aggregation of the cells = was=20 observed and most of the soma and neuronal processes showed = an=20 intense immunoreaction with anti-neuronal specific enolase. = (b) The=20 astrocytes were detected with anti-GFAP. The cells showed a = radial=20 morphology and represented 20% of the cultured cells. (c) = The soma=20 and cell processes of the neurons showed an intense staining = with=20 anti-SVCT2. (d=96e) Analysis of vitamin C transport in = primary=20 cultures of cerebellar rat neurons. (d) Uptake of = 100 μM AA in=20 the presence of NaCl or replacing NaCl with choline chloride = to=20 37=B0C. (e) Kinetic analysis of the AA transport. = Dose=96response of the=20 transport of AA to 37=B0C. (f) Eadie=96Hofstee plot for the = substrate=20 dependence of AA transport. Data represent the = mean =B1 SD=20 of two experiments. Scale bars: (a=96c) = 30 μm.=20

[Normal View=20 ]

We examined AA transport in rat neurons isolated = from=20 cerebellar cortex. The total uptake in 10 min was=20 270 pmol =D7 10⁶ cells (Fig. 5d).=20 Transport of AA by neurons was decreased by at least 70% in the = absence of=20 sodium and the total uptake was=20 92 pmol =D7 10⁶ cells in = 10 min=20 (Fig. 5d).=20 Dose=96response studies using 5 min assays revealed that the = transport=20 of AA in neurons is saturated at concentrations higher than = 400 μM=20 (Fig. 5e).=20 The Eadie=96Hofstee transformation showed apparent K_m of 20 and = 140 μM and a=20 V_max of 160 and=20 = 650 pmol =D7 10⁶cells/min,=20 respectively (Fig. 5f).

=20

=20

Neuroblastoma cell lines and human = tumors=20 express SVCT2

Most of the neurons isolated from brain cortex,=20 neuroblastoma (Neuro2a) and immortalized hippocampal neuronal = (HN33.11)=20 cell lines were positive to anti-βIII tubulin and anti-MAP2 = antisera (Fig. 6b,=20 e, and h and data not shown). We also analyzed the expression = of SVCT=20 isoform in these cells by immunohistochemistry using the specific = antibody=20 to SVCT2. Intense immunoreaction was detected in the soma and = processes of=20 the neurons, Neuro2a and HN33.11 cells (Fig. 6a,=20 d, and g). Additionally, the expression of SVCT2 was confirmed = by=20 using RT-PCR. A band of 339 bp was observed in mRNA samples = isolated=20 from brain cortex neurons, Neuro2a and HN33.11 cells (Fig. 6J,=20 lanes 2=964). To confirm that SVCT2 detected in neuroblastoma cell = lines is=20 also expressed in brain tumors we performed an immunohistochemical = analysis using neuroblastoma tumors isolated from human brain. A = weak but=20 consistent immunoreaction in the tumor cells was observed; = however, most=20 of the reaction was detected intracellularly (Fig. 7a=96c).=20 Additionally, we analyzed the expression of SVCT1 and the GLUT1 = (Fig. 7d=96i).=20 Our result only confirmed GLUT1 expression in neuroblastoma tumor = (Fig. 7g=96i).

=20

= Fig. 6=20 Immunohistochemical characterization and RT-PCR = analysis of SVCT2 expression in embryonic rat neurons, = Neuro2a, and=20 HN33.11 cell lines. An intense immunoreaction for tubulin = and SVCT2=20 was detected in 7-day cultured neurons (a=96c), = neuroblastoma Neuro2a=20 (d=96f) and HN33.11 (g=96i) cell lines. In neurons, the = immunoreaction=20 was concentrated in the soma and cell processes. However, in = immortalized cells the reaction was mainly observed = intracellularly.=20 (j) RT-PCR analysis for SVCT2 and β-actin expression. = The PCR=20 products were separated on 1.2% agarose gels and visualized = by=20 staining with ethidium bromide. Lane 1, DNA 100-mer size = standard.=20 The numbers on the left indicate the base pairs. Lane 2, = positive=20 controls using mRNA isolated from brain cortex neurons. Lane = 3,=20 RT-PCR products obtained using mRNA isolated from Neuro2a = cell=20 lines. Lane 4, RT-PCR products obtained using mRNA isolated = from=20 HN33.11 cell lines. Lane 5, reactions in which the cDNA = synthesis=20 step was performed in absence of the reverse transcriptase = (RT−)=20 using the mRNA isolated from brain cortex neurons. Scale = bars: (a=96i)=20 12 μm.

[Normal View=20 ]

=20

= Fig. 7=20 Immunohistochemical characterization of SVCTs = and GLUT1=20 expression in human neuroblastoma tumors. The = immunofluorescence=20 analysis was performed using secondary antibodies labeled = with Cy2=20 (green). The nucleus was stained with propidium iodide = (red).=20 Intense immunoreaction for SVCT2 and GLUT1 was detected in a = neuroblastoma tumor. SVCT2 was detected in the cellular = membrane=20 (a=96c), however; GLUT1 immunoreaction was observed = intracelullarly=20 (g=96i). SVCT1 was not detected in neuroblastoma tumor = cells. Scale=20 bars: 20 μm.

[Normal View=20 ]

=20

=20

Neuro2a and HN33.11 express only = SVCT2 but=20 demonstrate two different functional kinetic components involved = in AA=20 transport

We examined the AA transporter present in HN33.11 = and=20 Neuro2a cells. In HN33.11 cells the transport of AA was linear for = the=20 first 15 min (Fig. 8a).=20 The initial velocity of the AA uptake was=20 = 30 pmol =D7 10⁶cells/min. = The=20 total uptake in 15 min was=20 820 pmol =D7 10⁶ cells (Fig. 8a).=20 Transport of AA by neurons was decreased by at least 70% in the = absence of=20 sodium and the uptake reached a plateau of=20 180 pmol =D7 10⁶ cells in = 10 min=20 (Fig. 8a).=20 Similarly, in Neuro2a cells the transport of AA was linear for the = first=20 10 min and the initial velocity of the AA uptake was=20 = 25 pmol =D7 10⁶cells/min.=20 Transport of AA by Neuro2a cells was decreased by at least 90% in = the=20 absence of sodium and the uptake reached a plateau of=20 15 pmol =D7 10⁶ cells in = 3 min=20 (Fig. 8e).

=20

= Fig. 8 = Kinetic=20 analysis of vitamin C uptake in neuroblastoma cell lines. = Analysis=20 of vitamin C transport in Neuro2a and HN33.11 cell lines. (a = and e)=20 Time-course uptake of 100 μM AA in the presence of = NaCl or=20 replacing NaCl with choline chloride to 37=B0C.(b=96f)) = Time-course=20 uptake of 25, 50, and 100 μM AA in the presence of = NaCl to=20 37=B0C. (c and g) Kinetic analysis of the AA transport. = Dose=96response=20 of the transport of AA to 37=B0C. Eadie=96Hofstee plot for = the substrate=20 dependence of AA transport (d and h). Data represent the=20 mean =B1 SD of three experiments. =

[Normal=20 View ]

Next, we analyzed the uptake of vitamin C in = HN33.11 and=20 Neuro2a cells. Uptake proceeded in a linear fashion for at least=20 10 min at a substrate concentration of 25=96100 μM = (Fig. 8b=20 and f). A detailed dose=96response study showed that the = transport of AA=20 approached saturation at about 160 μM substrate (Fig. 8c=20 and g). Analysis of the transport data by the Eadie=96Hofstee = method=20 revealed the presence of two different functional components, = involved in=20 AA transport by HN33.11 and Neuro2a cells (Fig. 8d=20 and h). In HN33.11 cells, the higher affinity component had an = apparent transport K_m of=20 7 μM and a V_max of=20 25 pmol/min per 10⁶ cells, whereas = the lower=20 affinity component had apparent K_m and V_max values of = 86 μM and=20 60 pmol/min per 10⁶ cells, = respectively. In=20 Neuro2a cells, the higher affinity component had an apparent = transport=20 K_m of = 13 μM and a V_max of 40 pmol/min = per=20 10⁶ cells, whereas the lower affinity = component=20 had apparent K_m and = V_max values of = 105 μM and=20 110 pmol/min per 10⁶ cells,=20 respectively.

=20

=20

AA uptake in neurons and = neuroblastoma cells=20 is inhibited by flavonoids

When the cortical neurons were incubated in the = presence of=20 200 μM quercetin, AA transport was decreased by 80% (Fig. 9a,=20 bar 2). A similar effect was observed in cerebellar neurons (Fig. 9b,=20 bar 2). To further characterize the inhibitory action of different = flavonoid molecules on the AA uptake, we compared the quercetin = inhibitory=20 effect with the inhibition activity of phloretin. A similar = inhibition=20 effect (60=9670%) was observed when neurons isolated from brain = cortex and=20 cerebellum were incubated with phloretin (Fig. 9a=20 and b, bars 3). We did not observe any effect on AA transport = when=20 either cytochalasin B or E (GLUT inhibitors) was used (Fig. 9a=20 and b, bars 4 and 5). Similar results were observed in HN33.11 = and=20 Neuro2a cells (Fig. 9d=96f).=20 Dose-dependent inhibition analysis showed that quercetin inhibited = AA=20 transport in cortical neurons and Neuro2a cells with=20 IC₅₀ value of 30 μM and = 40 μM,=20 respectively (Fig. 9c=20 and f).

=20

= Fig. 9 = Acute=20 effect of inhibitors on the initial rates of AA uptake. = Brain and=20 cerebellar cortex neurons, Neuro2a, and HN33.11 cells lines = were=20 pre-treated with the different inhibitors for 10 min.=20 Time-course uptake of 100 μM ascorbic acid (AA) = replacing NaCl=20 with choline chloride (1), 200 μM quercetin (2), = 300 μM=20 phloretin (3), cytochalasin E (4), and cytochalasin B (5). = Semi-log=20 plot of the concentration dependence for inhibition of AA = transport=20 by quercetin in brain cortex neurons (C) or Neuro2a cell = lines (F).=20 Data represent the mean =B1 SD of two = experiments.=20

[Normal View=20 ]

=20

=20

Discussion

=

=

=

Using the techniques of RT-PCR, western blot, = in situ hybridization and immunohistochemical = analysis, our=20 data confirms the expression of SVCT2 transporter in developing = rat brain.=20 Furthermore, our functional study, using cultured neurons isolated = from=20 brain cortex and cerebellar cortex, demonstrated that SVCT2 is = localized=20 in the cellular membrane and is involved in the uptake of vitamin = C by the=20 cultured neurons. Surprisingly, we observed the presence of two = affinity=20 constants suggesting a regulatory mechanism associated with the = uptake of=20 AA in these cells. A similar observation was found in Neuro2a and = HN33.11=20 cells, demonstrating that this kinetic property is maintained in=20 immortalized neurons. An apparent K_m for AA transport in=20 neuroblastoma cells SH-SY5Y of 113 μM has been reported = (May=20 et al. 2006), which is comparable = to the=20 higher K_m observed = in our=20 studies in cultured neurons and neuroblastoma cell lines Neuro2a = and=20 HN33.11 (Castro=20 et al. 2001 and present data). A = similar=20 component has been observed in primary culture of cortical neurons = with an=20 apparent K_m for AA = transport=20 of 115 μM (Qiu=20 et al. 2007). These data indicate = that SVCT2=20 transporter (apparent K_m > 85 μ= M in=20 neurons and neuroblastoma cells) may be responsible for the = transport and=20 accumulation of near millimolar concentrations of AA inside these = cells=20 (Rice=20 2000). Our first kinetic analysis of AA uptake in mouse = cortical=20 neurons demonstrated the presence of two affinity constants, 8 and = 103 μM (Castro=20 et al. 2001). Similar data have = been=20 obtained in this study using rat neurons isolated from brain = cortex (K_m of 11 and = 92 μM) and=20 cerebellum (K_m of = 20 and=20 140 μM). Furthermore, we substantiated our data using = immortalized=20 neuronal cell lines Neuro2a (K_m of 13 and = 100 μM) and=20 HN33.11 (K_m of 7 = and=20 86 μM). These data raise the possibility that neurons = and=20 neuroblastoma cells may have specific kinetic regulatory = mechanisms to=20 transport vitamin C. For instance, the expression of very high = affinity AA=20 transporters may suggest a specific mechanism present in neurons, = and in=20 other cell types (neuroblastoma cells), to take up vitamin C under = pathological conditions, so that extracellular vitamin C can be = maintained=20 at a very low concentration (Chauhan=20 et al. 1987; Kallner=20 1987; Spielholz=20 et al. 1997). To understand the = kinetic=20 regulatory mechanism associated with the uptake of AA in neurons = and=20 neuroblastoma cells additional experiments are necessary. Two = binding=20 sites for AA on a single transporter (Hamill=20 et al. 1999) or a differential = interaction=20 with the sodium binding site (Godoy=20 et al. 2007) are possibilities; = however,=20 this needs to be experimentally demonstrated.

It has been demonstrated that flavonoids have a = novel=20 regulatory action on the uptake of vitamin C by sodium-dependent = AA=20 transporters. Flavonoids reversibly inhibit vitamin C transport in = SVCT1=20 transfected cells with IC₅₀ values of=20 10=9650 μM (Song=20 et al. 2002). Quercetin is a = reversible and=20 non-competitive inhibitor of AA transport, therefore, it has been=20 suggested that flavonoids may modulate vitamin C uptake in = different cells=20 that express SVCT1. However, there is no data associated to the = effect=20 produced by flavonoids in the uptake of vitamin C by SVCT2 = expressed in=20 neurons. In this work, we analyzed the effect of quercetin and = phloretin=20 on the uptake of AA in SVCT2 expressing neurons isolated from = brain and=20 cerebellar cortex, HN33.11 and Neuro2a neuroblastoma cell lines. = Our=20 results confirm that flavonoids modulate vitamin C transport in = SVCT2=20 expressing cells (Biondi=20 et al. 2007).

Ascorbic acid promotes osteogenic differentiation = in=20 mesenchymal stem cells (Jaiswal=20 et al. 1997), has been implicated = in=20 neuronal development and functional maturation (Qiu=20 et al. 2007), and possibly = performs a role=20 in the differentiation of embryonic cortical precursors into = neurons and=20 astrocytes (Lee=20 et al. 2003). Real-time PCR = analysis has=20 suggested that AA increases 1.5- to 2.5-fold the expression of = different=20 genes (NeuroD, Notch, BMP2, and BMP7) involved in neuronal and = glial=20 differentiation (Lee=20 et al. 2003). Thus, it is feasible = that AA=20 may play fundamental roles in glial and neuronal differentiation. = To=20 validate this hypothesis it is necessary to demonstrate that the = precursor=20 cells of the brain and, possibly, the migrating neurons and glial = cells=20 express vitamin C transporters associated with the uptake of this=20 nutrient.

Our in situ hybridization = and=20 immunohistochemical analysis strongly suggest that SVCT2 is = predominantly=20 expressed in the VZ and SVZ of the rat developing brain. = Additionally,=20 SVCT2 transporter is expressed in brain neuroepithelial cells and = choroid=20 plexus cells. Therefore, vitamin C present in the CSF may be = incorporated=20 into the precursor cells that are in contact with the ventricular = cavity.=20 These findings raise interesting questions concerning the role of = SVCT2=20 transporter and vitamin C in brain development.

The presence of SVCT2 in CNS precursors suggests = an early=20 role for vitamin C in the developing brain. It has been = demonstrated that=20 AA treatment promoted differentiation of dopaminergic and = serotoninergic=20 neurons (Yan=20 et al. 2001). AA treatment in = cultured=20 dopaminergic neurons was more effective during days 1=962 of = differentiation=20 than during days 3=965, indicating that AA affects an early step = of=20 dopaminergic neuronal differentiation (Yan=20 et al. 2001). Thus, AA may = regulate specific=20 genes associated with neuronal differentiation and thereby = contribute to=20 the fate of specific neurons.

We detected a high expression of the SVCT2 = transporter in=20 the ventricular and subventricular area of the embryonic brain = (where the=20 precursor cells are localized), which suggests that = neuroepithelial cells=20 may need vitamin C for proliferation and, possibly, for initial=20 differentiation. If so, to support these functions the = concentration of=20 vitamin C in the CSF need to be strictly regulated. Whole-body=20 radiography, at various times after ¹⁴C-AA = injection, shows that AA from blood penetrates the choroid plexus = first=20 and then enters into the CSF and finally the brain (Spector=20 and Lorenzo 1973; Rice=20 2000). It has been postulated that the choroid plexus cell is = the=20 place of AA entry into the CSF and is involved in the regulation = of=20 vitamin C concentration in the CSF. In adult brain it has been = documented=20 that the blood=96CSF=96brain barrier of the choroid plexus cells = accumulates=20 vitamin C in CSF 10-fold over the normal plasma level of = 57 μM (Rice=20 2000). The normal concentration of vitamin C in the CSF of = embryonic=20 brain has not been determined, however, it is possible that the = early=20 differentiation of choroid plexus cells induces a premature = formation of=20 the blood=96CSF barrier, which is involved in the concentration of = vitamin C=20 in the CSF. GLUT1 expression has been considered a marker of = blood=96CSF=20 barrier differentiation. We have demonstrated in human fetal brain = that=20 GLUT1 is expressed early in choroid plexus cells (Nualart=20 et al. 1999), this is consistent = with the=20 concept that blood=96CSF barrier appears to form early in the = fetal human=20 brain.

We detected a strong immunoreaction for SVCT2 in = rat choroid=20 plexus exactly in the proliferating area of the plexus that = corresponds to=20 the base of the structure. The differentiation of these cells = induced a=20 down-regulation of SVCT2 expression. However, SVCT2 was also = detected in=20 the cytoplasm of differentiated cells localized in the tip of the = plexus.=20 We were unable to observe a polarized localization of the SVCT2=20 transporter in the choroidal plexus cells; therefore, no = conclusion can be=20 drawn regarding the exact mechanism associated with transcellular=20 transport of vitamin C in embryonic plexus = cells.

=20

=20

=20

Acknowledgments

=

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This study was supported by Grants FONDECYT = 3020007 and=20 CONICYT-World Bank ACT-02 and ACI-12.

=20

=20

References

=

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