{\rtf\ansi\deff0\ansicpg1252{\fonttbl{\f0\froman times new roman;}{\f1\fswiss helvetica;}}{\colortbl\red0\green0\blue0;\red167\green169\blue172;\red211\green71\blue60;\red255\green255\blue255;\red0\green112\blue186;}\jexpand\viewkind1\viewscale100{\shp{\*\shpinst\shpleft10493\shptop493\shpright10666\shpbottom520\shpfhdr0\shpwr3\shpwrk0\shpfblwtxt1\shplid2028\shpz0\shpbxpage\shpbypage{\sp{\sn shapeType}{\sv 0}}{\sp{\sn fFlipH}{\sv 0}}{\sp{\sn fFlipV}{\sv 0}}{\sp{\sn rotation}{\sv 0}}{\sp{\sn geoRight}{\sv 173}}{\sp{\sn geoBottom}{\sv 27}}{\sp{\sn shapePath}{\sv 4}}{\sp{\sn pVerticies}{\sv 8;5;(0,0);(0,40);(186,40);(186,0);(0,0);}}{\sp{\sn pSegmentInfo}{\sv 2;12;16384;45824;1;45824;1;45824;1;45824;1;45824;24577;32768}}{\sp{\sn fFillOK}{\sv 1}}{\sp{\sn fFilled}{\sv 1}}{\sp{\sn fillColor}{\sv 11780847}}{\sp{\sn fLine}{\sv 0}}{\sp{\sn lineColor}{\sv 0}}{\sp{\sn lineType}{\sv 0}}{\sp{\sn fArrowheadsOK}{\sv 0}}{\sp{\sn fBehindDocument}{\sv 1}}{\sp{\sn fLayoutInCell}{\sv 1}}}}{\shp{\*\shpinst\shpleft10680\shptop493\shpright10866\shpbottom520\shpfhdr0\shpwr3\shpwrk0\shpfblwtxt1\shplid2029\shpz1\shpbxpage\shpbypage{\sp{\sn shapeType}{\sv 0}}{\sp{\sn fFlipH}{\sv 0}}{\sp{\sn fFlipV}{\sv 0}}{\sp{\sn rotation}{\sv 0}}{\sp{\sn geoRight}{\sv 186}}{\sp{\sn geoBottom}{\sv 27}}{\sp{\sn shapePath}{\sv 4}}{\sp{\sn pVerticies}{\sv 8;5;(0,0);(0,40);(186,40);(186,0);(0,0);}}{\sp{\sn pSegmentInfo}{\sv 2;12;16384;45824;1;45824;1;45824;1;45824;1;45824;24577;32768}}{\sp{\sn fFillOK}{\sv 1}}{\sp{\sn fFilled}{\sv 1}}{\sp{\sn fillColor}{\sv 4016340}}{\sp{\sn fLine}{\sv 0}}{\sp{\sn lineColor}{\sv 0}}{\sp{\sn lineType}{\sv 0}}{\sp{\sn fArrowheadsOK}{\sv 0}}{\sp{\sn fBehindDocument}{\sv 1}}{\sp{\sn fLayoutInCell}{\sv 1}}}}{\shp{\*\shpinst\shpleft10880\shptop493\shpright11066\shpbottom520\shpfhdr0\shpwr3\shpwrk0\shpfblwtxt1\shplid2030\shpz2\shpbxpage\shpbypage{\sp{\sn shapeType}{\sv 0}}{\sp{\sn fFlipH}{\sv 0}}{\sp{\sn fFlipV}{\sv 0}}{\sp{\sn rotation}{\sv 0}}{\sp{\sn geoRight}{\sv 186}}{\sp{\sn geoBottom}{\sv 27}}{\sp{\sn shapePath}{\sv 4}}{\sp{\sn pVerticies}{\sv 8;5;(0,0);(0,40);(186,40);(186,0);(0,0);}}{\sp{\sn pSegmentInfo}{\sv 2;12;16384;45824;1;45824;1;45824;1;45824;1;45824;24577;32768}}{\sp{\sn fFillOK}{\sv 1}}{\sp{\sn fFilled}{\sv 1}}{\sp{\sn fillColor}{\sv 11780847}}{\sp{\sn fLine}{\sv 0}}{\sp{\sn lineColor}{\sv 0}}{\sp{\sn lineType}{\sv 0}}{\sp{\sn fArrowheadsOK}{\sv 0}}{\sp{\sn fBehindDocument}{\sv 1}}{\sp{\sn fLayoutInCell}{\sv 1}}}}\paperw11880\paperh15560\margl666\margr133\margt573\margb66\pard\li9200\ri0\sl-186\slmult0 \fs16\cf1\f0\charscalex100 {Research Reports}\par\pard\li360\ri0\sl-213\slmult0 \fs18\cf0\par\pard\li360\ri0\sl-680\slmult0 \fs46\cf2\f0\charscalex100 {Bioluminescent CXCL12 fusion protein for cellular}\par\pard\li360\ri0\sl-546\slmult0 \fs46\cf2\f0\charscalex100 { studies of CXCR4 and CXCR7}\par\pard\li360\ri0\sl-346\slmult0 \fs24\cf0\f0\charscalex100 {Kathryn E. Luker}\fs16\cf0\f0\charscalex100 {1}\fs24\cf0\f0\charscalex100 {, Mudit Gupta}\fs16\cf0\f0\charscalex100 {1}\fs24\cf0\f0\charscalex100 {, and Gary D. Luker}\fs16\cf0\f0\charscalex100 {1, 2}\par\pard\li360\ri0\sl-333\slmult0 \fs16\cf0\f0\charscalex100 {1}\fs22\cf0\f0\charscalex100 {Center for Molecular Imaging, Department of Radiology, University of Michigan Medical School, Ann Arbor, MI, USA}\par\pard\li360\ri0\sl-333\slmult0 \fs24\cf0\f0\charscalex100 {and}{ }\fs16\cf0\f0\charscalex100 {2}\fs22\cf0\f0\charscalex100 {Department of Microbiology and Immunology, University of Michigan Medical School, Ann Arbor, MI, USA}\par\pard\li360\ri0\sl-266\slmult0 \fs12\cf0\f0\charscalex100 {BioTechniques}\fs12\cf0\f0\charscalex100 { 47:625-632 (July 2009) doi 10.2144/000113126 }\par\pard\li360\ri0\sl-226\slmult0 \fs12\cf0\f0\charscalex100 {Keywords: chemokine; bioluminescence; chemokine receptor;}{ }\fs16\cf0\f0\charscalex100 {Gaussia}\fs16\cf0\f0\charscalex100 { luciferase}\par\pard\li360\ri0\sl-213\slmult0 \fs18\cf0\par\pard\li360\ri0\sl-320\slmult0 \fs24\cf0\f0\charscalex100 {Chemokine CXCL12 and its two known receptors, CXCR4 and CXCR7, may play a role in diseases including}\par\pard\li360\ri0\sl-306\slmult0 \fs20\cf0\f0\charscalex100 {tumor growth and metastasis, atherosclerosis, and HIV infection. Therefore, these molecules may be promising tar}\fs20\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-306\slmult0 \fs20\cf0\f0\charscalex100 {gets for drug development. While studies of cell signaling and high-throughput screening for drug discovery increas}\fs20\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-306\slmult0 \fs20\cf0\f0\charscalex100 {ingly are based on luminescent assays because of their high sensitivity and signal-to-background ratio, there currently}\par\pard\li360\ri0\sl-306\slmult0 \fs22\cf0\f0\charscalex100 {is no bioluminescent assay for chemokine\uc1\u8211Xchemokine receptor binding. To develop a bioluminescent probe for}\par\pard\li360\ri0\sl-333\slmult0 \fs20\cf0\f0\charscalex100 {chemokine binding and cellular uptake, we fused CXCL12 to}{ }\fs22\cf0\f0\charscalex100 {Gaussia}\fs20\cf0\f0\charscalex100 { luciferase (GL), an ATP-independent enzyme}\par\pard\li360\ri0\sl-333\slmult0 \fs22\cf0\f0\charscalex100 {that is the smallest known luciferase. Fusing CXCL12 to}{ }\fs24\cf0\f0\charscalex100 {Gaussia}\fs22\cf0\f0\charscalex100 { luciferase (CXCL12-GL) did not alter the biolu}\fs22\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-306\slmult0 \fs20\cf0\f0\charscalex100 {minescence emission spectrum and only minimally affected enzyme function under varying conditions of pH, tem}\fs20\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-306\slmult0 \fs22\cf0\f0\charscalex100 {perature, and NaCl concentration. CXCL12-GL also activated CXCR4-dependent signaling to a comparable extent }\par\pard\li360\ri0\sl-306\slmult0 \fs22\cf0\f0\charscalex100 {as unfused CXCL12. Using multiwel plate assays, we established that CXCR7 increases cel -associated CXCL12}\par\pard\li360\ri0\sl-306\slmult0 \fs22\cf0\f0\charscalex100 {to a significantly greater extent than CXCR4. We also showed that CXCL12-GL can be used to quantify inhibition}\par\pard\li360\ri0\sl-306\slmult0 \fs22\cf0\f0\charscalex100 {of chemokine receptor binding by compounds that specifical y target CXCR7. These data validate CXCL12-GL as}\par\pard\li360\ri0\sl-306\slmult0 \fs22\cf0\f0\charscalex100 {a bioluminescent probe to investigate molecular functions of CXCR4 and CXCR7 and screen for compounds that}\par\pard\li360\ri0\sl-306\slmult0 \fs24\cf0\f0\charscalex100 {modulate ligand-receptor binding.}\par\pard\sect\sectd\sbknone\cols3\colno1\colw3813\colsr-0\colno2\colw3466\colsr-0\colno3\colw3800\pard\li360\ri0\sl-213\slmult0 \fs18\cf0\par\pard\li360\ri0\sl-213\slmult0 \fs18\cf0\par\pard\li360\ri0\sl-426\slmult0 \fs28\cf2\f0\charscalex100 {Introduction}\par\pard\li360\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {Chemokines and chemokine receptors }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {originally were identified as regulators of }\par\pard\li360\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {immune cell trafficking in normal physi-}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {ology and inflammation. More recently, }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {this family of proteins has been shown to }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {regulate diseases including cancer, athero}\fs18\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {sclerosis, autoimmune diseases, and neuro}\fs18\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {degenerative processes (1,2\uc1\u8211X4). As a result,}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {chemokine receptors have emerged as new}\par\pard\li360\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {therapeutic targets, emphasizing the need }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {to develop reagents for quantitative, high-}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {throughput screening assays of chemokine}\par\pard\li360\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {binding and inhibition.}\par\pard\li600\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {Conventional assays use radiolabeled }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {chemokines to quantify binding to specific}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {receptors and identify inhibitors of ligand-}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {receptor interactions. While radioligand }\par\pard\li360\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {binding studies have been used successfully }\par\pard\li360\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {for these purposes, such assays generate}\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {radioactive waste that is cumbersome to }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {dispose. As alternatives to radioactivity, }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {investigators increasingly are using fluores}\fs18\cf0\f0\charscalex100 {-}\par\pard\li360\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {cence or bioluminescence for cell-based}\par\pard\li360\ri0\sl-253\slmult0 \fs14\cf0\f0\charscalex100 {assays in basic research and high-throughput }\par\pard\li360\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {screening. CXCL12 and other chemokines }\par\pard\li360\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {have been labeled with fluorescent dyes or }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {proteins to detect ligand-receptor binding }\par\pard\li360\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {in intact cells (5). For example, Hatse et al.}\par\pard\li360\ri0\sl-226\slmult0 \fs18\cf1\f0\charscalex100 {Vol. 47 | No. 1 | 2009}\par\column\pard\li0\ri0\sl-213\slmult0 \fs18\cf0\par\pard\li0\ri0\sl-453\slmult0 \fs18\cf0\f0\charscalex100 {fused CXCL12 to AlexaFluor 647 and used }\par\pard\li0\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {this reagent to detect binding and inhibition }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {of CXCL12-CXCR4 interactions by }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {flow cytometry (6). Similarly, binding of }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {CXCL12 to receptor CXCR7 has been }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {analyzed with a genetic fusion of CXCL12}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {to green fluorescent protein (GFP) (7).}\par\pard\li240\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {Whilethesestudiesshowthatfluorescent }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {chemokines provide a viable replacement for }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {radiolabeled chemokine binding assays in }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {some experimental settings, biolumines}\fs20\cf0\f0\charscalex100 {-}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {cence assays have not been developed for }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {chemokine-chemokine receptor binding }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {studies. Relative to fluorescence-based }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {techniques, bioluminescence assays with }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {luciferase enzymes offer several potential }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {advantages for quantifying accumulation}\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {of chemokines in intact cells (8). First, }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {bioluminescence assays have substantially }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {lower background signal than fluores}\fs20\cf0\f0\charscalex100 {-}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {cence, enhancing signal-to-noise ratios for }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {chemokine binding. Improved sensitivity }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {is particularly important for measuring }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {ligand binding to chemokine and other }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {seven-transmembrane receptors because}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {these proteins may be expressed at modest}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {levels in cells (9). Second, bioluminescent }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {end points have a greater dynamic range of}\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {linear signal response than corresponding }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {fluorescent probes. Finally, bioluminescence }\par\pard\li1093\ri0\sl-453\slmult0 \fs36\cf1\f0\charscalex100 {625}\par\column\pard\li0\ri0\sl-213\slmult0 \fs18\cf0\par\pard\li0\ri0\sl-453\slmult0 \fs18\cf0\f0\charscalex100 {assays are affected less than fluorescence}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {by colors and dyes in compounds and cell }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {culture media. These advantages account}\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {for improved performance of biolumines}\fs20\cf0\f0\charscalex100 {-}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {cence relative to fluorescence in multiwell }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {cell culture formats and highlight the need }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {for a bioluminescent assay of chemokine-}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {chemokine receptor binding (10).}\par\pard\li240\ri0\sl-253\slmult0 \fs16\cf0\f0\charscalex100 {To develop a bioluminescent assay for }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {binding of chemokines to receptors, we}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {focused on chemokine CXCL12 and its two }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {known receptors, CXCR4 and CXCR7. }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {There is particular interest in developing }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {specific inhibitors of CXCL12 binding }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {to these two receptors both as chemical}\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {probes for investigating specific mecha}\fs20\cf0\f0\charscalex100 {-}\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {nisms of action and as potential therapeutic }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {agents for treating diseases including cancer }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {and HIV (11\uc1\u8211X13). We fused CXCL12 to }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {the humanized form of luciferase from }\par\pard\li0\ri0\sl-280\slmult0 \fs18\cf0\f0\charscalex100 {Gaussia princeps}\fs20\cf0\f0\charscalex100 {.}{ }\fs20\cf0\f0\charscalex100 {Gaussia}\fs18\cf0\f0\charscalex100 { luciferase (GL) is}\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {the smallest known luciferase, containing }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {only 185 amino acids. In cell-based assays, }\par\pard\li0\ri0\sl-266\slmult0 \fs20\cf0\f0\charscalex100 {GL is}{ }\fs20\cf0\f0\charscalex100 {\uc1\u8764X}\fs20\cf0\f0\charscalex100 {1000 brighter than firefly and }\par\pard\li0\ri0\sl-280\slmult0 \fs20\cf0\f0\charscalex100 {Renil a}\fs18\cf0\f0\charscalex100 { luciferases (two other enzymes used }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {commonly for bioluminescent assays), which }\par\pard\li0\ri0\sl-253\slmult0 \fs18\cf0\f0\charscalex100 {increases the dynamic range for detection }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {and quantification (14). Bioluminescence }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {from GL is also independent of ATP, }\par\pard\li0\ri0\sl-253\slmult0 \fs20\cf0\f0\charscalex100 {allowing the enzyme to function in both }\par\pard\li1453\ri0\sl-226\slmult0 \fs16\cf1\f0\charscalex100 {www.BioTechniques.com}\par\pard\sect\sectd\sbknone\pard\sect\sectd\sbkpage{\shp{\*\shpinst\shpleft813\shptop493\shpright1000\shpbottom520\shpfhdr0\shpwr3\shpwrk0\shpfblwtxt1\shplid2031\shpz0\shpbxpage\shpbypage{\sp{\sn shapeType}{\sv 0}}{\sp{\sn fFlipH}{\sv 0}}{\sp{\sn fFlipV}{\sv 0}}{\sp{\sn rotation}{\sv 0}}{\sp{\sn geoRight}{\sv 187}}{\sp{\sn geoBottom}{\sv 27}}{\sp{\sn shapePath}{\sv 4}}{\sp{\sn pVerticies}{\sv 8;5;(0,0);(0,40);(186,40);(186,0);(0,0);}}{\sp{\sn pSegmentInfo}{\sv 2;12;16384;45824;1;45824;1;45824;1;45824;1;45824;24577;32768}}{\sp{\sn fFillOK}{\sv 1}}{\sp{\sn fFilled}{\sv 1}}{\sp{\sn fillColor}{\sv 11780847}}{\sp{\sn fLine}{\sv 0}}{\sp{\sn lineColor}{\sv 0}}{\sp{\sn lineType}{\sv 0}}{\sp{\sn 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