Activity-dependent = immobilization of mitochondria: the role of Miro

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General Commentary

Activity-dependent immobilization of=20 mitochondria: the role of Miro

F. Anne=20 Stephenson ^*

School of Pharmacy, University of London, UK

A commentary on

Miro1 is a calcium sensor for glutamate = receptor-dependent localization of mitochondria at synapses

by MacAskill, A. F., Rinholm, J. E.,=20 Twelvetrees, A. E., Arancibia-Carcamo, L., Muir, J., Fransson, A., = Aspenstrom,=20 P., Attwell, D., and Kittler, J. T. (2009) Neuron 61,=20 541=96555.

It is well established that in neurones, = at any one=20 time, in the range of 13=9630% of the mitochondrial population are = mobile moving=20 in both anterograde and retrograde directions utilising motor proteins = and the=20 microtubule network. It is also known that the mobility of mitochondria = is=20 affected by neuronal activity; the empirical observation being that = mitochondria=20 are immobilized by a Ca²⁺-dependent mechanism in response to = release=20 of neurotransmitters or post-synaptic receptor activation presumably to = satisfy=20 the high energy demands of synaptic transmission. The molecular players = involved=20 in these transitions and their modes of action are now beginning to be=20 unravelled. Two recent papers shed new light on mechanisms of = mitochondrial=20 dynamics (MacAskill=20 et al., 2009; Wang=20 and Schwarz, 2009). This article is principally a commentary on the = MacAskill=20 et al. (2009) paper but the findings reported therein are discussed = with=20 respect to those reported concurrently by Wang=20 and Schwarz (2009).

The focus of MacAskill=20 et al. (2009) is the mitochondrial protein, Miro1, and its role on = the=20 mobility of dendritic mitochondria in primary cultures of hippocampal = neurones.=20 Miro1 (and Miro2) is an atypical RhoGTPase that is found in the outer = membrane=20 of all mitochondria. It has a short C-terminal domain that is located = inside=20 mitochondria, one transmembrane domain and within the N-terminal region = which=20 protrudes from the mitochondria into the cell cytoplasm, it has two = GTPase=20 domains, I and II, which are separated by two EF-hand calcium binding = domains,=20 EF-hands I and II. First, MacAskill=20 et al. (2009) showed that over-expression of green fluorescent = protein=20 (GFP)-Miro1 resulted in ∼ 20% increase of mobile mitochondria with = no change in=20 neither their density nor average velocity. Conversely, shRNAi = knock-down of=20 Miro1 resulted in ∼ 10% decrease in the percentage of mobile = mitochondria.=20 Perfusion of GFP-Miro1 transfected neuronal cultures with glutamate and = glycine=20 to activate glutamate receptors or activation of L-type Ca²⁺ = channels=20 in the presence of extracellular Ca²⁺ resulted in the almost = complete=20 cessation of mitochondrial movement. Expression of a form of GFP-Miro in = which=20 the two EF-hand calcium binding domains were inactivated by mutation = i.e.,=20 GFP-Miro1 ΔEF, did not change basal mitochondrial dendritic = transport but=20 importantly, glutamate-induced arrest did not occur. Co-cultures were = used to=20 demonstrate that in the presence of perfused glutamate, immobilized = mitochondria=20 were accumulated at synapses, i.e. that there is an activity-dependent=20 positioning of mitochondria. Finally, rather than perfusing with = glutamate,=20 local stimulation of neurones with glutamate delivered via a patch = pipette=20 resulted in the halting of mitochondrial movement within a 15 =B5m = region for 150=20 s demonstrating that synaptic activation results in the recruitment of = passing=20 mitochondria to synapses.

This series of elegant experiments = utilizing=20 molecular and physiological approaches clearly demonstrates the pivotal = role of=20 Miro1 and its Ca²⁺ binding domain in mitochondrial transport = and=20 mitochondrial docking at synapses. Wang=20 and Schwarz (2009) attained a similar conclusion however the actual=20 molecular mechanism by which this occurs is a matter of dispute between = the=20 different groups. It is known that anterograde mitochondrial transport = is=20 mediated via interaction with kinesin motor proteins. Prior to the MacAskill=20 et al. (2009) publication, experimental evidence suggested a role = for the=20 Milton/TRAK family of kinesin adaptor proteins in mediating this = association (Glater=20 et al., 2006; Smith=20 et al., 2006). Milton/TRAKs bind to the cargo binding domain of = kinesin and=20 have been shown to recruit kinesin to mitochondria i.e. mitochondrial = (i.e.=20 Miro1)/kinesin association was indirect. In the MacAskill study, this = dogma is=20 challenged. Firstly, they showed that introduction of a kinesin=20 function-blocking antibody into neurones resulted in a decrease in the=20 percentage of mobile mitochondria proving the engagement of motor = proteins.=20 Kinesin/Miro1 interactions were then studied by in vitro pull = down=20 assays. In the absence of Ca²⁺, a direct association between = the=20 kinesin heavy chains, KIF5A, 5B and 5C, and Miro1 was detected. In the = presence=20 of 2 mM Ca²⁺ however, the association of Miro1 with these = motors was=20 significantly reduced. Miro1 ΔEF was also shown to bind kinesin and = this=20 association was, as may have been predicted, = Ca²⁺-independent.=20 Co-immunoprecipitations and pull downs from extracts of rat brain = ratified these=20 in vitro findings. Thus it was concluded that the presence of=20 Ca²⁺ inhibits the Miro1/kinesin protein=96 protein = interaction hence=20 the motor is dissociated from mitochondria yielding arrested = movement.

As implicated above, the study of Wang=20 and Schwarz (2009) also addressed the mechanism of = Ca²⁺-mediated=20 regulation of mitochondrial motility but their findings contrasted with = those=20 reported by MacAskill=20 et al. (2009). Here, the focus was mitochondrial movement in axons = of=20 primary cultures of hippocampal neurones. Wang=20 and Schwarz (2009) found that at least when over-expressed, kinesin = is=20 present on all mitochondria. In the presence of Milton (the Drosophila = homologue=20 of TRAK), and Miro, kinesin became highly concentrated on mitochondria = providing=20 evidence that Milton (and presumably TRAKs) are required for the = recruitment of=20 kinesin to neuronal mitochondria. Similarly to MacAskill=20 et al. (2009), axonal mitochondrial transport was measured following = expression of Miro1. Elevation of intracellular Ca²⁺ by = calcimycin=20 addition to neuronal cultures resulted in reduced bidirectional = mitochondrial=20 transport. This decreased mobility was not apparent following expression = of=20 Miro1 ΔEF. Under high Ca²⁺ conditions, there was no = evidence for a=20 release of kinesin from the mitochondria and, intriguingly, = immunoprecipitations=20 now revealed an association between kinesin (the motor domain) and = Miro1. This=20 protein=96protein interaction was concomitant with an observed reduced = binding of=20 kinesin to microtubules. The interpretation of these findings by the = authors is=20 that in the absence of Ca²⁺, Milton/TRAKs are required to = recruit=20 kinesin to mitochondria for their transport. In the presence of elevated = Ca²⁺ however, the motor domain of kinesin binds directly to = Miro1 in=20 preference to microtubules thus the motor becomes detached from the = transporting=20 network resulting in the arrest of mitochondrial movement.

In summary, both papers and also a paper = from Saotome=20 et al. (2008), are compatible in terms of Miro1 being a key player = in=20 mitochondrial transport and activity-dependent docking at synapses. = However, the=20 findings of the two groups result in the proposal of two disparate = mechanisms by=20 which this occurs. How can this be reconciled? The biochemical studies = of MacAskill=20 et al. (2009) focussed on in vitro interactions between = Miro and=20 kinesin heavy chains in the presence and absence of Ca²⁺ but, = in the=20 absence of Milton/TRAKs. Wang=20 and Schwarz (2009) included Miro, kinesin heavy chain and Milton for = co-immunoprecipitation assays from transfected cells, a limitation here = being=20 that the three proteins are over-expressed. Despite these caveats, it = may be=20 speculated however that Ca²⁺ may be important for the = stabilization=20 of the Miro, kinesin, Milton/TRAK) complex. In the presence of high=20 Ca²⁺ and in the absence of Milton, the kinesin/Miro1 = interaction is=20 not stabile and it is dissociated from mitochondrial membranes. In the = presence=20 of Milton/TRAK, even in high Ca²⁺, kinesin remains associated = with=20 mitochondria but the mode of interaction is altered i.e. rather than = interacting=20 via the Milton/TRAK-Miro linkage, there is a switch such that the = kinesin motor=20 domain does not bind to microtubules but rather associates directly with = Miro1.=20 This suggests that both proposed mechanisms may operate; in the presence = of=20 Milton/TRAKs, regulation by Ca²⁺ may be a transient response = whereas=20 when Milton/TRAK is absent, the effect of high Ca²⁺ is more = sustained=20 due to dissociation from mitochondria. So a key question is what = controls the=20 formation of kinesin, Milton/TRAK complexes? Further, Wang=20 and Schwarz (2009) found that manipulating Miro1 expression also had = effects=20 on retrograde mitochondrial transport implying that Miro1 associates = with=20 dynein. Saotome=20 et al. (2008) reported that the effects of Miro are not limited to=20 transport. They found that Miro plays a role in mitochondrial = fusion=96fission=20 switching and Ca²⁺ signalling. There has been a surge of = activity in=20 this field due to proteins involved in mitochondrial transport being = recently=20 identified. Further advances that may in the long term impact on = understanding=20 the pathogenesis of neurodegenerative diseases in which defective = trafficking is=20 implicated are awaited with keen anticipation.

Acknowledgments

Work in the author=92s laboratory is = supported by the=20 BBSRC (UK) and the Alzheimer=92s Research Trust (UK).

References

Glater, E. E., Megeath, L. J., Stowers, R. = S., and=20 Schwarz, T. L. (2006). Axonal transport of mitochondria requires Milton = to=20 recruit kinesin heavy chain and is light chain independent. J. Cell=20 Biol. 173, 545=96557.

Pubmed=20 Abstract Pubmed=20 Full Text CrossRef=20 Full Text

MacAskill, A. F., Rinholm, J. E., = Twelvetrees, A. E.,=20 Arancibia-Carcamo, L., Muir, J., Fransson, A., Aspenstrom, P., Attwell, = D., and=20 Kittler, J. T. (2009). Miro1 is a calcium sensor for glutamate=20 receptor-dependent localization of mitochondria at synapses. = Neuron 61,=20 541=96555.

Pubmed=20 Abstract Pubmed=20 Full Text CrossRef Full = Text=20

Saotome, M., Safiulina, D., Szabadkai, G., = Das, S.,=20 Fransson, A., Aspenstrom, P., Rizzuto, R., and Hjnoczky, G. (2008).=20 Bidirectional Ca²⁺-dependent control of mitochondrial = dynamics by the=20 Miro GTPase. Proc. Natl. Acad. Sci. U.S.A. 105, = 20728=9620733.

Pubmed=20 Abstract Pubmed=20 Full Text CrossRef=20 Full Text

Smith, M. J., Pozo, K., Brickley, K., and = Stephenson,=20 F. A. (2006). Mapping the GRIF-1 binding domain of the kinesin, KIF5C,=20 substantiates a role for GRIF-1 as an adaptor protein in the anterograde = trafficking of cargoes. J. Biol. Chem. 281, = 27216=9627228.

Pubmed=20 Abstract Pubmed=20 Full Text CrossRef=20 Full Text

Wang, X. and Schwarz, T. L. (2009). The = mechanism of=20 Ca²⁺-dependent regulation of kinesin-mediated mitochondrial = motility.=20 Cell 136, 163=96174.

Pubmed=20 Abstract Pubmed=20 Full Text CrossRef Full = Text=20

Citation: Stephenson F (2010) = Activity-dependent=20 immobilization of mitochondria: the role of Miro. Front. Mol. = Neurosci.=20 3:9. 1-2. doi:10.3389/fnmol.2010.00009

Received: = 20 January=20 2010; Accepted: 17 March 2010; Published online: 13 April=20 2010.

Edited=20 by:

Morgan Sheng, Massachusetts Institute of = Technology,=20 USA

Reviewed=20 by:

Yi-Ping Hsueh, Institute of Molecular = Biology, Academia=20 Sinica, Taiwan
Casper Hoogenraad, Erasmus Medical Center,=20 Netherlands

Copyright: =A9 2010 Stephenson. This is = an=20 open-access article subject to an exclusive license agreement between = the=20 authors and the Frontiers Research Foundation, which permits = unrestricted use,=20 distribution, and reproduction in any medium, provided the original = authors and=20 source are credited.

*Correspondence: F. Anne Stephenson, = School of=20 Pharmacy, University of London, 29/39 Brunswick Square, London WC1N 1AX, = UK.=20 e-mail: anne.stephenson@pharmacy.ac.uk

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Dec 12th, 08" (v1.01): Fixed Shadow issue when multiple LIs within = the same UL (level) contain sub menus: = http://www.dynamicdrive.com/forums/showthread.php?t=3D39177&highlight=3Ds= mooth //** Feb 11th, 09" (v1.02): The currently active main menu item (LI A) = now gets a CSS class of ".selected", including sub menu items. //** May 1st, 09" (v1.3): //** 1) Now supports vertical (side bar) menu mode- set "orientation" to = 'v' //** 2) In IE6, shadows are now always disabled //** July 27th, 09" (v1.31): Fixed bug so shadows can be disabled if = desired. var ddsmoothmenu=3D{ //Specify full URL to down and right arrow images (23 is padding-right = added to top level LIs with drop downs): arrowimages: {down:['downarrowclass', 'down.gif', 23], = right:['rightarrowclass', '/images/sub0.gif', '/images/sub1.gif']}, transition: {overtime:300, outtime:300}, //duration of slide in/ out = animation, in milliseconds shadow: {enable:false, offsetx:5, offsety:5}, ///////Stop configuring beyond here/////////////////////////// detectwebkit: = navigator.userAgent.toLowerCase().indexOf("applewebkit")!=3D-1, //detect = WebKit browsers (Safari, Chrome etc) detectie6: document.all && !window.XMLHttpRequest, getajaxmenu:function($, setting){ //function to fetch external page = containing the panel DIVs var $menucontainer=3D$('#'+setting.contentsource[0]) //reference empty = div on page that will hold menu $menucontainer.html("Loading Menu...") $.ajax({ url: setting.contentsource[1], //path to external menu file async: true, error:function(ajaxrequest){ $menucontainer.html('Error fetching content. Server Response: = '+ajaxrequest.responseText) }, success:function(content){ $menucontainer.html(content) ddsmoothmenu.buildmenu($, setting) } }) }, buildmenu:function($, setting){ var smoothmenu=3Dddsmoothmenu var $mainmenu=3D$("#"+setting.mainmenuid+">ul") //reference main menu = UL $mainmenu.parent().get(0).className=3Dsetting.classname || = "ddsmoothmenu" //var firstLevelMenu =3D $(".ddsmoothmenu-v > *")[0]; //$(".ulLeft .liFirst>a").css("background", "#fff"); $(".ulLeft .liFirst>a").addClass("firstLevelMenu"); $(".ulLeftGrey .liFirst>a").addClass("firstLevelMenuGrey"); //$(".ulLeft .on").addClass("on"); =09 =09 var $headers=3D$mainmenu.find("ul").parent() $headers.hover( function(e){ $(this).children('a:eq(0)').addClass('selected') }, function(e){ $(this).children('a:eq(0)').removeClass('selected') } ) $headers.each(function(i){ //loop through each LI header var $curobj=3D$(this).css({zIndex: 100-i}) //reference current LI = header var $subul=3D$(this).find('ul:eq(0)').css({display:'block'}) this._dimensions=3D{w:this.offsetWidth, h:this.offsetHeight, = subulw:$subul.outerWidth(), subulh:$subul.outerHeight()} this.istopheader=3D$curobj.parents("ul").length=3D=3D1? true : false = //is top level header? $subul.css({top:this.istopheader && setting.orientation!=3D'v'? = this._dimensions.h+"px" : 0}) $curobj.children("a:eq(0)").css(this.istopheader? {paddingRight: = smoothmenu.arrowimages.down[2]} : {}).append( //add arrow images '

' ) if (smoothmenu.shadow.disable){ = this._shadowoffset=3D{x:(this.istopheader?$subul.offset().left+smoothmenu= .shadow.offsetx : this._dimensions.w), y:(this.istopheader? = $subul.offset().top+smoothmenu.shadow.offsety : $curobj.position().top)} = //store this shadow's offsets if (this.istopheader) $parentshadow=3D$(document.body) else{ var $parentLi=3D$curobj.parents("li:eq(0)") $parentshadow=3D$parentLi.get(0).$shadow } this.$shadow=3D$('

').prependTo($parentshadow).css({left:this._shadowoffset.x+'= px', top:this._shadowoffset.y+'px'}) //insert shadow DIV and set it to = parent node for the next shadow div } $curobj.hover( =20 function(e){ var $targetul=3D$(this).children("ul:eq(0)") = $($targetul).parent(":first").children(":first").addClass("menuMouseOver"= ); = $($targetul).parent(":first").children(":first").html($($targetul).parent= (":first").children(":first").html().replace(smoothmenu.arrowimages.right= [1], smoothmenu.arrowimages.right[2])); this._offsets=3D{left:$(this).offset().left, = top:$(this).offset().top} var menuleft=3Dthis.istopheader && setting.orientation!=3D'v'? 0 : = this._dimensions.w = menuleft=3D(this._offsets.left+menuleft+this._dimensions.subulw>$(window)= .width())? (this.istopheader && setting.orientation!=3D'v'? = -this._dimensions.subulw+this._dimensions.w : -this._dimensions.w) : = menuleft //calculate this sub menu's offsets from its parent if ($targetul.queue().length<=3D1){ //if 1 or less queued animations $targetul.css({left:menuleft+"px", = width:this._dimensions.subulw+'px'}).animate({height:'show',opacity:'show= '}, ddsmoothmenu.transition.overtime) if (smoothmenu.shadow.enable){ var shadowleft=3Dthis.istopheader? = $targetul.offset().left+ddsmoothmenu.shadow.offsetx : menuleft var = shadowtop=3Dthis.istopheader?$targetul.offset().top+smoothmenu.shadow.off= sety : this._shadowoffset.y if (!this.istopheader && ddsmoothmenu.detectwebkit){ //in WebKit = browsers, restore shadow's opacity to full this.$shadow.css({opacity:1}) } this.$shadow.css({overflow:'', width:this._dimensions.subulw+'px', = left:shadowleft+'px', = top:shadowtop+'px'}).animate({height:this._dimensions.subulh+'px'}, = ddsmoothmenu.transition.overtime) } } }, function(e){ var $targetul=3D$(this).children("ul:eq(0)") = $($targetul).parent(":first").children(":first").removeClass("menuMouseOv= er"); 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Content-Location: http://owa.frontiersin.org/public/main.php?owa_view=base.jsLogLib // // Open Web Analytics - An Open Source Web Analytics Framework // // Copyright 2006 Peter Adams. All rights reserved. // // Licensed under GPL v2.0 http://www.gnu.org/copyleft/gpl.html // // Unless required by applicable law or agreed to in writing, software // distributed under the License is distributed on an "AS IS" BASIS, // WITHOUT WARRANTIES OR CONDITIONS OF ANY KIND, either express or = implied. // See the License for the specific language governing permissions and // limitations under the License. // // $Id$ // /**=0A= *=0A= * URL encode / decode=0A= * http://www.webtoolkit.info/=0A= *=0A= **/=0A= =0A= var Url =3D {=0A= =0A= // public method for url encoding=0A= encode : function (string) {=0A= return escape(this._utf8_encode(string));=0A= },=0A= =0A= // public method for url decoding=0A= decode : function (string) {=0A= return this._utf8_decode(unescape(string));=0A= },=0A= =0A= // private method for UTF-8 encoding=0A= _utf8_encode : function (string) {=0A= string =3D string.replace(/\r\n/g,"\n");=0A= var utftext =3D "";=0A= =0A= for (var n =3D 0; n < string.length; n++) {=0A= =0A= var c =3D string.charCodeAt(n);=0A= =0A= if (c < 128) {=0A= utftext +=3D String.fromCharCode(c);=0A= }=0A= else if((c > 127) && (c < 2048)) {=0A= utftext +=3D String.fromCharCode((c >> 6) | 192);=0A= utftext +=3D String.fromCharCode((c & 63) | 128);=0A= }=0A= else {=0A= utftext +=3D String.fromCharCode((c >> 12) | 224);=0A= utftext +=3D String.fromCharCode(((c >> 6) & 63) | 128);=0A= utftext +=3D String.fromCharCode((c & 63) | 128);=0A= }=0A= =0A= }=0A= =0A= return utftext;=0A= },=0A= =0A= // private method for UTF-8 decoding=0A= _utf8_decode : function (utftext) {=0A= var string =3D "";=0A= var i =3D 0;=0A= var c =3D c1 =3D c2 =3D 0;=0A= =0A= while ( i < utftext.length ) {=0A= =0A= c =3D utftext.charCodeAt(i);=0A= =0A= if (c < 128) {=0A= string +=3D String.fromCharCode(c);=0A= i++;=0A= }=0A= else if((c > 191) && (c < 224)) {=0A= c2 =3D utftext.charCodeAt(i+1);=0A= string +=3D String.fromCharCode(((c & 31) << 6) | (c2 & = 63));=0A= i +=3D 2;=0A= }=0A= else {=0A= c2 =3D utftext.charCodeAt(i+1);=0A= c3 =3D utftext.charCodeAt(i+2);=0A= string +=3D String.fromCharCode(((c & 15) << 12) | ((c2 = & 63) << 6) | (c3 & 63));=0A= i +=3D 3;=0A= }=0A= =0A= }=0A= =0A= return string;=0A= }=0A= =0A= } /** * Javascript Tracking Library *=20 * @author Peter Adams * @copyright Copyright © 2006 Peter Adams = * @license http://www.gnu.org/copyleft/gpl.html GPL v2.0 * @category owa * @package owa * @version $Revision$ =20 * @since owa 1.0.0 */ var OWA =3D {} =09 OWA.log =3D function() { =09 this.id =3D ''; } OWA.log.prototype =3D { =20 // private method for issuing logging request _makeAjaxRequest : function (properties) { =09 var bug var get var init =20 url =3D this._assembleRequestUrl(properties); =20 if (window.XMLHttpRequest){ =09 // If IE7, Mozilla, Safari, etc: Use native object var ajax =3D new XMLHttpRequest() =09 }=20 =09 else { =09 if (window.ActiveXObject){ =09 // ...otherwise, use the ActiveX control for IE5.x and IE6 var ajax =3D new ActiveXObject("Microsoft.XMLHTTP");=20 } =09 } =20 =09 ajax.open("GET", url, false);=20 ajax.send(null); =09 // Uninitialize variable. init =3D null; =09 return; }, // private method for issuing logging request _makeRequest : function (properties) { =09 var bug var url =20 url =3D this._assembleRequestUrl(properties); bug =3D "

"; =20 document.write(bug); =09 return; }, =20 _assembleRequestUrl : function(properties) { =20 var get var log_url =09 get =3D ''; =09 log_url =3D 'http://owa.frontiersin.org/public/log.php?'; =09 //assemble query string for(param in properties) { // print out the params =20 value =3D ''; =09 if (typeof properties[param] !=3D 'undefined') { =09 value =3D Url.encode(this._base64_encode(properties[param]+'')); =09 } else { =09 value =3D ''; =09 } =20 get =3D get + "owa_" + param + "=3D" + value + "&"; } =09 // add some radomness for cache busting return log_url + get + Math.round(100*Math.random()); =20 }, =20 _base64_encode : function(decStr) { =20 var base64s =3D = 'ABCDEFGHIJKLMNOPQRSTUVWXYZabcdefghijklmnopqrstuvwxyz0123456789+/'; var bits; var dual; var i =3D 0; var encOut =3D ''; =09 while(decStr.length >=3D i + 3) { bits =3D (decStr.charCodeAt(i++) & 0xff) <<16 | (decStr.charCodeAt(i++) & 0xff) <<8 | decStr.charCodeAt(i++) & 0xff; =09 encOut +=3D base64s.charAt((bits & 0x00fc0000) >>18) + base64s.charAt((bits & 0x0003f000) >>12) + base64s.charAt((bits & 0x00000fc0) >> 6) + base64s.charAt((bits & 0x0000003f)); } =09 if(decStr.length -i > 0 && decStr.length -i < 3) { dual =3D Boolean(decStr.length -i -1); =09 bits =3D ((decStr.charCodeAt(i++) & 0xff) <<16) | (dual ? (decStr.charCodeAt(i) & 0xff) <<8 : 0); =09 encOut +=3D base64s.charAt((bits & 0x00fc0000) >>18) + base64s.charAt((bits & 0x0003f000) >>12) + (dual ? base64s.charAt((bits & 0x00000fc0) >>6) : '=3D') = + '=3D'; } =09 return(encOut); } } // OWA Page View object ///////////////////////////////////// OWA.pageView =3D function(caller_params) { this.properties =3D new Object(); =09 for(param in caller_params) { // print out the params =20 this.properties[param] =3D caller_params[param]; =09 } this._setProperties(); =09 return; }; OWA.pageView.prototype =3D { // public method for setting logging request properties _setProperties : function () { =20 this.properties["event"] =3D "base.page_request";=09 this.properties["action"] =3D "base.processRequest"; =20 if (typeof this.properties["page_uri"] =3D=3D 'undefined') { this.properties["page_url"] =3D document.URL; } if (typeof this.properties["page_title"] =3D=3D 'undefined') { this.properties["page_title"] =3D document.title; } =09 if (typeof this.properties["referer"] =3D=3D 'undefined') { this.properties["referer"] =3D document.referrer; } =20 return; }, =20 log : function() { =20 logger =3D new OWA.log; return logger._makeRequest(this.properties); =20 } } // OWA click object ///////////////////////////////////////// /** * Constructor of the click object * * @param e Event Object */ OWA.click =3D function(caller_params) { this.properties =3D new Object(); =09 for(param in caller_params) { // print out the params =20 this.properties[param] =3D caller_params[param]; =09 } =20 this.e =3D ''; =09 return; }; OWA.click.prototype =3D { properties : new Object, =09 e : '', =09 init: 0, =09 =09 /** * Sets all properties of the click object * * @param e Event Object */ setProperties : function(e) { =09 this.e =3D e; this.properties["event"] =3D "base.click"; this.properties["action"] =3D "base.processEvent"; this._setTarget(); this._setTagName(); this._setCoords(); this.properties["dom_element_name"] =3D this.targ.name; this.properties["dom_element_value"] =3D this.targ.value; this.properties["dom_element_id"] =3D this.targ.id; this.properties["page_url"] =3D window.location.href; this.init =3D 1; =20 return true; }, =09 log : function() { =09 //alert(this.properties["site_id"]); //alert("hello from click.log"); =09 if (this.init =3D=3D 1) { =09 var logger =09 logger =3D new OWA.log; logger._makeAjaxRequest(this.properties); } =09 this.init =3D 0; return; =09 }, =09 /** * Sets coordinates of where in the browser the user clicked * */ _setCoords : function() { =09 var windowWidth =3D window.innerWidth ? window.innerWidth : = document.body.offsetWidth; var windowHeight =3D window.innerHeight ? window.innerHeight : = document.body.offsetHeight; =09 if( typeof( this.e.pageX ) =3D=3D 'number' ) { =09 this.properties["click_x"] =3D this.e.pageX + ''; this.properties["click_y"] =3D this.e.pageY + ''; } else { =20 this.properties["click_x"] =3D this.e.clientX + ''; this.properties["click_y"] =3D this.e.clientY + ''; } =09 this.properties["dom_element_x"] =3D this._findPosX(this.targ) + = ''; this.properties["dom_element_y"] =3D this._findPosY(this.targ) + = ''; this.properties["page_width"] =3D windowWidth + ''; this.properties["page_height"] =3D windowHeight + ''; =09 return; }, =09 /** * Sets the X coordinate of where in the browser the user clicked * */ _findPosX : function(obj) { =09 var curleft =3D 0; if (obj.offsetParent) { while (obj.offsetParent) { curleft +=3D obj.offsetLeft obj =3D obj.offsetParent; } } else if (obj.x) curleft +=3D obj.x; return curleft; }, =09 /** * Sets the Y coordinates of where in the browser the user clicked * */ _findPosY : function(obj) { =09 var curtop =3D 0; if (obj.offsetParent) { while (obj.offsetParent) { curtop +=3D obj.offsetTop obj =3D obj.offsetParent; } } else if (obj.y) curtop +=3D obj.y; return curtop; }, =09 /** * Sets the HTML element that actually generated the event * */ _setTarget : function() { =09 // Determine the actual html element that generated the event //if (this.e.target) { // this.targ =3D this.e.target; =20 //} else if (this.e.srcElement) { // this.targ =3D this.e.srcElement; // } =20 this.targ =3D this.e.target || this.e.srcElement; =20 if (this.targ.nodeType =3D=3D 3) { // defeat Safari bug this.targ =3D target.parentNode; } =20 return; }, =09 /** * Sets the tag name of html eleemnt that generated the event */ _setTagName : function() { =20 // Set properties of the owa_click object. this.properties["dom_element_tag"] =3D this.targ.tagName; =20 if (this.properties["dom_element_tag"] =3D=3D "A") { =20 if (this.targ.textContent !=3D undefined) { this.properties["dom_element_text"] =3D = this.targ.textContent; } else { this.properties["dom_element_text"] =3D = this.targ.innerText; } =20 this.properties["target_url"] =3D this.targ.href; =20 } else if (this.properties["dom_element_tag"] =3D=3D "INPUT") { =20 this.properties["dom_element_text"] =3D this.targ.value; } =20 else if (this.properties["html_element_tag"] =3D=3D "IMG") { =20 this.properties["target_url"] =3D this.targ.parentNode.href; this.properties["dom_element_text"] =3D this.targ.alt; } =20 else { =20 this.properties["target_url"] =3D this.targ.parentNode.href; =09 if (this.targ.textContent !=3D undefined) { this.properties["html_element_text"] =3D = this.targ.textContent; } else { this.properties["html_element_text"] =3D = this.targ.innerText; } } =09 return; }=09 } =0A= //=0A= // Open Web Analytics - An Open Source Web Analytics Framework=0A= //=0A= // Copyright 2006 Peter Adams. All rights reserved.=0A= //=0A= // Licensed under GPL v2.0 http://www.gnu.org/copyleft/gpl.html=0A= //=0A= // Unless required by applicable law or agreed to in writing, software=0A= // distributed under the License is distributed on an "AS IS" BASIS,=0A= // WITHOUT WARRANTIES OR CONDITIONS OF ANY KIND, either express or = implied.=0A= // See the License for the specific language governing permissions and=0A= // limitations under the License.=0A= //=0A= // $Id$=0A= //=0A= =0A= =0A= /**=0A= * Logs Page View=0A= *=0A= * Takes owa_param object which is defined by the logging tag.=0A= *=0A= * @param owa_params Object=0A= */=0A= var owa_pv =3D new OWA.pageView(owa_params);=0A= owa_pv.log();=0A= =0A= =0A= =0A= =0A= /**=0A= * Logs Click=0A= *=0A= * Takes owa_param object which is defined by the logging tag.=0A= *=0A= * @param owa_params Object=0A= */=0A= //Log Clicks=0A= var owa_click =3D new OWA.click(owa_params);=0A= =0A= =0A= /**=0A= * Helper function for setting properties on the click object=0A= *=0A= * Takes a DOM event object=0A= *=0A= * @param e Object=0A= */=0A= function owa_setClick(e) {=0A= =0A= // hack for IE7=0A= e =3D e || window.event;=0A= =0A= owa_click.setProperties(e);=0A= return;=0A= =0A= }=0A= =0A= /**=0A= * Helper Function for calling the log method on the lick object=0A= *=0A= */=0A= function owa_logClick() {=0A= =0A= owa_click.log();=0A= return;=0A= }=0A= =0A= // Registers the handler for the on.click event so that click properties = can be set=0A= document.onclick =3D owa_setClick;=0A= =0A= // Registers the handler for the beforenavigate event so that the click = can be logged=0A= =0A= if(window.addEventListener) {=0A= =0A= window.addEventListener('beforeunload', owa_logClick, false);=0A= =0A= } else if(window.attachEvent) {=0A= =0A= window.attachEvent('beforeunload', owa_logClick);=0A= =0A= }=0A= =0A= ------=_NextPart_000_0000_01CB1778.65168480--

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